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Institute
- Theodor-Boveri-Institut für Biowissenschaften (21)
- Institut für Physikalische und Theoretische Chemie (6)
- Physikalisches Institut (6)
- Institut für Virologie und Immunbiologie (5)
- Institut für Anorganische Chemie (4)
- Institut für Informatik (4)
- Institut für Molekulare Infektionsbiologie (4)
- Institut für Mathematik (3)
- Institut für Organische Chemie (3)
- Institut für Pharmakologie und Toxikologie (3)
RS1, a gene product of RSC1A1, is critically involved in cell density-dependent transcriptional down-regulation of SGLT1 in LLC-PK1 cells and in the post-transcriptional down-regulation of SGLT1 in small intestine. RS1 inhibits the release of SGLT1 containing vesicles from the trans-Golgi network and migrates into the nucleus where it inhibits transcription of SGLT1. In the present work we identified a novel 21 amino acids-long nonconventional nuclear localization sequence (RS1 NLS) in porcine RS1 (pRS1) that is necessary and sufficient for nuclear targeting of pRS1. RS1 NLS is framed by two consensus sequences for phosphorylation which are responsible for confluence-dependent regulation of RS1 NLS: a casein kinase 2 (CK2) site in position 348 and a protein kinase C (PKC) site in position 370. Confluence-dependent nuclear targeting was observed with amino acids 342-374 (R-NLS-Reg). Mutation analysis suggested that nuclear targeting is blocked by phosphorylation of serine 370 (PKC) and that phosphorylation of serine 348 (CK2) prevents phosphorylation of serine 370. Because CK2 is down-regulated and PKC is up-regulated during confluence of LLC-PK1 cells, our data suggest that nuclear localization coordinates cell density-dependent changes in transcriptional and post-transcriptional inhibition of SGLT1 expression.
1. Oviposition site selection is crucial for the reproductive success of herbivorous insects. According to the preference–performance hypothesis, females should oviposit on host plants that enhance the performance of their offspring. More specifically, the plant vigour hypothesis predicts that females should prefer large and vigorously growing host plants for oviposition and that larvae should perform best on these plants. 2. The present study examined whether females of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) prefer to oviposit on large host plant individuals of the meadow clary and whether large host plants are of higher nutritional quality than small host plants. Subsequently, it was tested whether the female preference correlates with offspring performance and survival. 3. In the field, females preferred large host plant individuals for oviposition and host plant quality, i.e. leaf nitrogen content, was significantly higher in leaves of large than of small host plants. 4. In the laboratory, larval development time was shorter on leaves of large host plant individuals than on small host plant individuals, but this could not be shown in the field. 5. However, a predator-exclusion experiment in the field resulted in a higher survival of larvae on large host plants than on small host plants when all predators had free access to the plants. On caged host plants there was no difference in survival of larvae between plant size categories. 6. It is concluded that females of C. canaliculata select oviposition sites that enhance both performance and survival of their offspring, which meets the predictions of the plant vigour hypothesis.
The present thesis reports about vibrational and quantum chemical investigations on model systems undergoing photochemical processes and pharmaceutically active compounds, respectively. Infrared (IR) and Raman spectroscopy were applied for the characterization of the ground state molecular structure. Moreover, resonance Raman (RR) spectra contain additional information about the resonantly enhanced excited state molecular structure. A quantitative resonance Raman intensity analysis in conjunction with the simultaneous simulation of the absorption spectra by means of time-dependent propagation methods was accomplished in order to extract valuable information about the excited state molecular structures of the investigated systems. Surface enhanced Raman scattering (SERS) allows one to determine the interaction and adsorption site of active agents on a metal substrate. Furthermore, density functional theory (DFT) and potential energy distribution (PED) calculations were carried out for an exact assignment of the vibrational spectra. Complete active space self consistent field (CASSCF) and configuration interaction (CI) calculations for some model systems were also performed to assess the experimental results on the excited state potential surfaces. The fundamentals of resonance Raman spectroscopy are treated in detail, describing the physical processes and emphasizing the theoretical methodologies which allow one to obtain the information about the resonantly excited state via an RR intensity analysis. The Brownian oscillator model to determine the solvent reorganization energy is briefly presented. Furthermore, the SERS enhancement mechanisms and selection rules to determine the orientation of the molecules adsorbed on the metal substrate are discussed. The Hartree-Fock approach to calculate the ground state geometry is expatiated, and the basic characteristics of the CI and CASSCF calculations are specified. The chapter ends with a short description of the DFT calculations. Chapter 4 deals with the investigation of the excited state intramolecular proton transfer of the model system, 1-hydroxy-2-acetonaphthone (HAN). The vibrations showing the highest displacement parameters correspond to stretching and in-plane deformation modes of the naphthalene ring and the conjugated carbonyl group, while the OH stretching mode exhibits no observable enhancement. The cooperative effect of the skeletal vibrations reduces the distance between the carbonyl and hydroxyl oxygen atoms in accordance with a general electron density redistribution. Hence, the leading force in the proton transfer process is the increase in electron density on the carbonyl group and the decrease of the negative charge on the hydroxyl oxygen. In chapter 5 the structural and vibrational characteristics of the organic mixed valence system N,N,N’,N’-tetraphenylphenylenediamine radical cation (1+) are discussed. The resonance Raman measurements showed that at least eight vibrational modes are strongly coupled to the optical charge transfer process in (1+). These Franck-Condon active modes were assigned to symmetric vibrations. The most enhanced band corresponds to the symmetric stretching mode along the N-phenylene-N unit and exhibits the largest vibrational reorganization energy. Nevertheless, symmetric stretching modes of the phenylene and phenyl units as well as deformation modes are also coupled to the electronic process. The total vibrational reorganization energy of these symmetrical modes is dominant, while the solvent induced broadening and reorganization energy are found to be small. Hence, (1+) adopts a symmetrical delocalized Robin-Day Class III structure in the ground state. Chapter 6 reports about a vibrational spectroscopic investigation of a model organic photorefractive thiophene derivative, 2-(N,N-diethylamino)-5-(2’,2’-dicyanovinyl)-thiophene. The geometry of the first excited state were optimized and the FC parameters were calculated using the configuration interaction with single excitations method. These calculations show that the contribution of the zwitterionic structure to the excited state is significantly higher than in the ground state. The resonance Raman spectra indicate that several stretching modes along the bonds connecting the donor and acceptor moieties as well as the S-C stretching vibrations are enhanced. Chapter 7 presents the vibrational analysis of an aziridinyl tripeptide, a cysteine protease inhibitor active drug. The vibrational analysis reveals stronger H-bonding of the aziridine NH unit in the solid state of the aziridinyl tripeptide than in the liquid electrophilic building block, indicating medium strong intermolecular H-bond interactions in the crystal unit. The amide hydrogen atoms of the aziridinyl tripeptide are involved in weaker H-bonds than in an epoxide analogon. Furthermore, the characteristic vibrational modes of the peptide backbone were discussed. Chapter 8 reports on the adsorption mechanism of two related anti-leukemia active agents, 6-mercaptopurine (6MP) and 6-mercaptopurine-ribose (6MPR) on a silver colloid. Both molecules adsorb through the N1 and possibly S atom on the metal surface under basic conditions. The SERS spectra recorded for acidic pH values showed that the ribose derivative exhibits a different adsorption behavior compared to the free base. 6MP probably adsorbs on the silver sol through the N9 and N3 atoms, while 6MPR interacts with the surface via the N7 and probably S atoms. Around critical biological concentrations and pH values i.e. at low concentrations and almost neutral condition (pH 7-9), 6MPR interacts with the substrate through both N7 and N1 atoms, possibly forming two differently adsorbed species, while for 6MP only the species adsorbed via N1 was evidenced.
We examined the regulation of NFATc1 in different lymphomas and observed an inversed correlation between the methylation status and expression of NFATc1. Our data demonstrate that aberrant DNA methylation associated with chromatin remodeling within nfatc1 locus is a major mechanism for the repression of NFATc1 expression, suggesting that the DNA methylation-mediated transcriptional silencing of NFATc1 may be a critical event in the tumorogenesis of ALCLs and cHLs. Furthermore, the DNA methylation of human nfatc1 promoter region could be used as a novel biomarker of tumor progression. Our results indicate a close link between the loss of immunoreceptor signaling and NFATc1 expression in human lymphomas. For both ALCLs and cHLs, defects in immunoreceptor signaling have been described which result in a loss of receptor-mediated gene expression programs (Schwering et al., 2003; Bonzheim et al., 2004; Marafioti et al., 2004). In T cells, one indicator gene of these programs appears to be the nfatc1 gene whose expression is controlled by TCR signals (Chuvpilo et al., 2002a). In contrast, in T cells NFATc1 expression is unaffected by TCR signals, and NFATc2 was found to be expressed at normal levels in ALCLs and cHLs (L.K., unpubl. data). Moreover, the activity of NF-kappaB factors which can bind to certain NFAT binding sites and share a distantly-related DNA binding domain with NFATs is strongly elevated in cHL cells (Bargou et al., 1997; Hinz et al., 2001; Hinz et al., 2002) suggesting that NFATs and NF-kappaBs exert very different effects on generation and maintenance of Hodgkin’s lymhomas. However, it should be mentioned that in Burkitt’s and further B cell lymphomas in which NFATc1 proteins are strongly expressed and controlled by receptor signals (Kondo et al., 2003), they could exert a promoting function in tumor development. The genes of p53 family members p63 and p73 are prominent examples for mammalian genes whose products can act both as oncoproteins and tumor suppressor genes (Hibi et al., 2000; Stiewe and Putzer, 2002), and it is likely that more genes exist which encode both tumor suppressors and oncoproteins. It remains to be shown whether the nfatc1 gene is one of them.
An exhaustive discussion of constraint qualifications (CQ) and stationarity concepts for mathematical programs with equilibrium constraints (MPEC) is presented. It is demonstrated that all but the weakest CQ, Guignard CQ, are too strong for a discussion of MPECs. Therefore, MPEC variants of all the standard CQs are introduced and investigated. A strongly stationary point (which is simply a KKT-point) is seen to be a necessary first order optimality condition only under the strongest CQs, MPEC-LICQ, MPEC-SMFCQ and Guignard CQ. Therefore a whole set of KKT-type conditions is investigated. A simple approach is given to acquire A-stationarity to be a necessary first order condition under MPEC-Guiganrd CQ. Finally, a whole chapter is devoted to investigating M-stationary, among the strongest stationarity concepts, second only to strong stationarity. It is shown to be a necessary first order condition under MPEC-Guignard CQ, the weakest known CQ for MPECs.
The analysis of real data by means of statistical methods with the aid of a software package common in industry and administration usually is not an integral part of mathematics studies, but it will certainly be part of a future professional work. The present book links up elements from time series analysis with a selection of statistical procedures used in general practice including the statistical software package SAS Statistical Analysis System). Consequently this book addresses students of statistics as well as students of other branches such as economics, demography and engineering, where lectures on statistics belong to their academic training. But it is also intended for the practician who, beyond the use of statistical tools, is interested in their mathematical background. Numerous problems illustrate the applicability of the presented statistical procedures, where SAS gives the solutions. The programs used are explicitly listed and explained. No previous experience is expected neither in SAS nor in a special computer system so that a short training period is guaranteed. This book is meant for a two semester course (lecture, seminar or practical training) where the first two chapters can be dealt with in the first semester. They provide the principal components of the analysis of a time series in the time domain. Chapters 3, 4 and 5 deal with its analysis in the frequency domain and can be worked through in the second term. In order to understand the mathematical background some terms are useful such as convergence in distribution, stochastic convergence, maximum likelihood estimator as well as a basic knowledge of the test theory, so that work on the book can start after an introductory lecture on stochastics. Each chapter includes exercises. An exhaustive treatment is recommended. This book is consecutively subdivided in a statistical part and an SAS-specific part. For better clearness the SAS-specific part, including the diagrams generated with SAS, always starts with a computer symbol, representing the beginning of a session at the computer, and ends with a printer symbol for the end of this session. This book is an open source project under the GNU Free Documentation License.
The plant cuticle as an interface between the plant interior and the adjoining atmosphere plays an important role in any interaction between the plant and its environment. Transport processes across the cuticles were the object of countless research since many decades. However, bulk of the work done was focused on transport of lipophilic molecules. It is highly plausible to examine the penetration of lipophilic compounds, since the cuticle is dominated by lipophilic compartments itself, and the most crop protection agents have lipophilic character. As a result of this research, cuticular transport of lipophilic compounds is relatively well understood. Since several years, examinations were expanded on transport of hydrophilic molecules. In the present study, a direct comparison was made between transport properties of lipophilic and hydrophilic compounds, which allows an objective assessment of the mechanism governing their penetration. The results of this present study debunked the existence of two different pathways across isolated cuticles of Hedera helix (English ivy), a lipophilic and a hydrophilic pathway. This finding was supported by examinations regarding to accelerator and temperature effects on the mobility of both pathways, because the hydrophilic path is insensitive to them - in contrary to the lipophilic one. The lipophilic pathway is rigorously restricted to lipophilic molecules and the hydrophilic pathway is only accessible for hydrophilic molecules. Uncharged hydrophilic compounds can cross the cuticle even the molecules are of relatively large dimensions. In contrast to that, dissociable compounds with a molar volume higher than 110 cm³ mol-1 are excluded from cuticular penetration. Differences in the mobility of uncharged and dissociable molecules might be a hint towards the chemical nature of the polar pathways. It is assumed, that both, cellulose and pectin fibrils, traverse the cuticle which are originated from the epidermal cell wall. While uncharged carbohydrates might be able to penetrate across a pathway made up of cellulose and pectin, dissociated amino acids might be restricted to the cellulose path. This could be a plausible explanation for the higher mobility and the higher cuticle/water partition coefficients of the carbohydrates compared with the amino acids. A hydrophilic pathway was found with isolated grapevine cuticles, too. The apparent size selectivity of the hydrophilic pathway implies transport via narrow pores. From the present data, a mean pore radius of 0.31 nm (H. helix) or rather 0.34 nm (V. vinifera) was calculated. The absolute number of pores per cm² is 1.1 x 109 for H. helix and 3.3 x 109 for V. vinifera cuticles. This finding and the enlarged pore size distribution of grapevine cuticles might be an explanation for the transport of uncharged and dissociable hydrophilic compounds of higher molar volume like paraquat dichloride - in contrast to ivy membranes Wax extraction of ivy membranes uncovers additional pores, which explains the increased mobilities of the hydrophilic compounds across dewaxed membranes. From these extensive measurements it is very conspicuous, that the bulk of cuticular water transpiration occurs via the polar pathway. Since the work was focused on cuticular penetration of primary metabolites like amino acids and carbohydrates, a mechanistic explanation of leaching processes is obtained, simultaneously. In cuticular research, an inconsistent terminology regarding the transport path of the hydrophilic compounds was used. The term ‘hydrophilic pathway’ is definitely correct, since it makes no statement with regard to the shape of this path. In contrast to that, the terms ‘polar pore’ or ‘aqueous pore’ could imply that there is a tube or rather a water-filled tube traversing the cuticle. However - at this point of time – the imagination about the shape of this path is a pathway across interfibrilar gaps within polysaccharide strains. The proposed diameter of these interfibrilar gaps fits very well to the diameter determined in this study. Therefore, the imagination of a pore is not unfounded, but it is a very narrow pore, definitely. Additionally, this pathway is a very straight pathway which corresponds to this simplified imagination. An expanded study was done with paraquat dichloride, which was applied as aqueous droplets on grapevine cuticles. It is assumed that these model membranes reflect transport properties which are very close to that of relevant crops and weeds. The predominating parameter for paraquat penetration is the moisture, either originated from a relative humidity of at least 75% or provided by added chemicals. There is a tendency for good suitability of hygroscopic additives. Increased paraquat penetration was also obtained by raised concentrations and removal of the cuticular waxes.
This thesis contributes to the field of silicon chemistry, with a special emphasis on the chemistry of penta- and hexacoordinate silicon.The spirocyclic zwitterionic Lambda5Si-silicates 1–6 with a (2,2,6,6-tetramethylpiperidinio)- methyl group and two identical bidentate chelate ligands derived from glycine, (S)-alanine, (S)-phenylalanine, (S)-valine, (S)-tert-leucine, or (S)-proline bound to the silicon(IV) coordination center were synthesized and structurally characterized for the first time.The hitherto unknown spirocyclic zwitterionic Lambda5Si-silicates 7–12 with an (ammonio)- methyl group and two identical bidentate chelate ligands derived from (S)-lactic acid, (S)-3- phenyllactic acid, or (S)-mandelic acid were synthesized and structurally characterized in the solid state (elemental analyses (C, H, N), crystal structure analyses, 15N and 29Si VACP/MAS solid-state NMR experiments) and in solution (except 10; 1H, 13C, and 29Si NMR experiments)The spirocyclic zwitterionic Lambda5Si-silicates 13, 15, and 16 with an (ammonio)methyl group and two bidentate meso-oxolane-3,4-diolato(2–) ligands bound to the silicon(IV) coordination center were synthesized for the first time. The already existent compound 14 was resynthesized in order to perform a crystal structure analysis. All compounds were characterized by elemental analyses (C, H, N), 29Si VACP/MAS solid-state NMR experiments, and solution NMR studies (1H, 13C, 15N, and 29Si NMR experiments), and compounds 14–16 were additionally studied by single-crystal X-ray diffraction.The already existent zwitterionic Lambda5Si-silicate 17 was synthesized by new methods, including a remarkable Si–C cleavage reaction with benzoin. To investigate the dynamic behavior of the known zwitterionic Lambda5Si-silicate 18 in solution, VT 1H NMR experiments in CD2Cl2 were performed in the temperature range –100 °C to 23 °C.The hexacoordinate silicon compounds 19–22 containing multidentate ligands derived from citric acid or (S)-malic acid were synthesized for the first time. The anionic Lambda6Si-silicates 19–22 were structurally characterized in the solid state by single-crystal X-ray diffraction and VACP/MAS NMR spectroscopy (13C, 15N, 29Si). Upon dissolution in water at 20 °C, spontaneous hydrolysis of the Lambda6Si-silicate anions was observed.
Diese Arbeit wurde durch Experimente zur Potential- und Stromverteilung in Quanten-Hall- Systemen motiviert, die in den letzten Jahren in der Abteilung von Klitzing am MPI für Festkörperforschung durchgeführt wurden und ergaben, dass elektrostatische Abschirmungseffekte in zweidimensionalen Elektronensystemen (2DES), die den ganzzahligen Quanten-Hall-Effekt (QHE) zeigen, sehr wichtig für das Verständnis der Stromverteilung innerhalb der Probe und der extremen Genauigkeit der gemessenen quantisierten Werte des Hall-Widerstands sind. Daraus ergab sich für die hier vorgelegte Arbeit das folgende Programm. Zunächst wird, nach einem einleitenden Kapitel, in Kapitel 2 der Formalismus vorgestellt, mit dem in den späteren Kapiteln Elektronendichten und elektrostatische Potentiale, die z.B. das 2DES auf eine Probe mit Streifengeometrie eingrenzen, selbstkonsistent berechnet werden. Diese Selbstkonsistenz besteht aus zwei Teilen. Erstens wird, bei vorgegebenem Potential, die Elektronendichte berechnet. Zweitens wird aus vorgegebener Ladungsverteilung, bestehend aus (positiven) Hintergrundladungen und der (im ersten Schritt berechneten) Elektronenladungsdichte, und geeigneten Randbedingungen (konstantes Potential auf metallischen Gates) durch Lösen der Poisson-Gleichung das elektrostatische Potential berechnet. Wenn wir im ersten Schritt, unter Berücksichtigung der Fermi-Dirac-Statistik, die Elektronendichte quantenmechanisch aus den Energieeigenfunktionen und -werten berechnen, erhalten wir die Hartree-Näherung, die die Dichte als nichtlokales Funktional des Potentials liefert. Wenn man die Ausdehnung der Wellenfunktionen auf der Längenskala, auf der sich das Potential typischerweise ändert, vernachlässigen kann, so vereinfacht sich die Hartree-Näherung zur Thomas- Fermi-Näherung, die einen lokalen Zusammenhang zwischen Elektronendichte und Potential beschreibt. Die meisten der konkreten Rechnungen wurden im Rahmen dieser selbstkonsistenten Thomas-Fermi-Poisson-Näherung durchgeführt. Im Kapitel 3 wird allgemein das Abschirmverhalten eines 2DES im hohen Magnetfeld untersucht. Wir betrachten die Antwort auf eine harmonische Potentialmodulation im unbegrenzten 2DES und in streifenförmig begrenzten Systemen mit zwei unterschiedlichen Arten von Randbedingungen. Bei tiefen Temperaturen und hohen Magnetfeldern finden wir extrem nichtlineare Abschirmung. Im unbegrenzten 2DES charakterisieren wir die Abschirmung, indem wir die gesamte Variation des selbstkonsistent berechneten Potentials als Funktion der Amplitude des aufgeprägten cosinus-Potentials berechnen. Bei festem Magnetfeld ergeben sich so Stufenfunktionen, deren Gestalt stark vom Füllfaktor der Landau-Niveaus im homogenen Zustand ohne aufgeprägtes Potential abhängt (siehe Abbildungen 3.2- 3.6). Vielleicht noch unerwartetere Kurven ergeben sich, wenn man bei festem Modulationspotential die Varianz des selbstkonsistenten Potentials gegen das Magnetfeld B aufträgt (Abb. 3.9). Die Resultate lassen sich aber leicht verstehen und (bei Temperatur T = 0) in einem einfachen Schema (Abb. 3.7) zusammenfassen. Als ordnendes Prinzip stellt sich heraus, dass sich stets Zustände einstellen, in denen die Elektronendichte möglichst wenig von der bei verschwindendem Magnetfeld abweicht. Wenn die Zyklotronenergie groß gegen die thermische Energie kBT ist, erfordert das, dass in den großen Bereichen, in denen die Dichte variiert, ein Landau-Niveau unmittelbar an dem, im Gleichgewicht konstanten, elektrochemischen Potential liegen muss (En, “pinning”). Man nennt diese Bereiche kompressibel. In den kompressiblen Bereichen können Elektronen leicht umverteilt werden, d.h. die Dichte ist leicht veränderbar und in diesen Bereichen gibt es extrem effektive Abschirmung. Existieren kompressible Bereiche mit unterschiedlichen Landau-Niveaus (En) am elektrochemischen Potential, z.B. bei großer Modulation oder weil die Dichte zum Probenrand hin abnimmt, so gibt es zwischen benachbarten kompressiblen Bereichen mit unterschiedlichen Landau-Quantenzahlen n “inkompressible” Bereiche, in denen zwischen zwei Landau-Niveaus liegt. Dort sind alle Landau-Niveaus unterhalb von besetzt, die oberhalb leer. Folglich ist dort der Füllfaktor ganzzahlig und die Dichte konstant. Das Wechselspiel zwischen kompressiblen und inkompressiblen Bereichen bestimmt das Abschirmverhalten. Randeffekte erweisen sich nur in solchen Magnetfeldintervallen als wichtig für die Abschirmung im Inneren einer streifenförmigen Probe, in denen (schon ohne aufgeprägte Modulation) in der Probenmitte ein neuer inkompressibler Streifen entsteht. Im Kapitel 4 wird die Rolle der inkompressiblen Streifen in einer idealisierten, streifenförmigen Hall-Probe untersucht. Mithilfe einer lokalen Version des Ohmschen Gesetzes berechnen wir bei vorgegebenen Gesamtstrom die Stromdichte und das nun ortsabhängige elektrochemische Potential, dessen Gradient die Stromdichte treibt. Für den lokalen Leitfähigkeitstensor nehmen wir ein für homogenes 2DES berechnetes Resultat und ersetzen den Füllfaktor jeweils durch den lokalen Wert. Dadurch ergibt sich, dass bei Existenz inkompressibler Streifen der gesamte Strom auf diese Streifen eingeschränkt ist, in denen die Komponenten des spezifischen Widerstands die Werte des freien, idealen 2DES haben, also verschwindenden longitudinalen und quantisierten Hall-Widerstand. Aus Hartree-Rechnungen zeigen wir, dass es inkompressible Streifen nur in Magnetfeldintervallen endlicher Breite (um ganzzahlige Füllfaktoren) gibt und dass in der Nähe von Füllfaktor 4 es nur inkompressible Streifen mit dem lokalen Füll-faktor \nu(x) = 4 gibt, aber nicht solche mit \nu(x) = 2, in Gegensatz zu dem Ergebnis der Thomas-Fermi-Poisson-Näherung, die hier nicht gültig ist. Um diese Unzulänglichkeit der Thomas-Fermi-Poisson-Näherung und Artefakte des strikt lokalen Modells zu beheben, führen wir die Rechnungen mit einem (auf der Skala des mittleren Elektronenabstands) gemittelten Leitfähigkeitstensors aus. Damit erhalten wir, im Rahmen einer Linear-Response-Rechnung, sehr schöne Übereinstimmung mit den Potentialmessungen, die diese Dissertation motivierten, einen kausalen Zusammenhang zwischen der Existenz inkompressibler Streifen und der Existenz von Plateaus im QHE, und ein Verständnis der extremen Genauigkeit, mit der die quantisierten Widerstandswerte reproduziert werden können, unabhängig von Probenmaterial und -geometrie. Im Kapitel 5 untersuchen wir das Zufallspotential, in dem sich die Elektronen bewegen. Wir gehen davon aus, dass sich hinter einer undotierten Schicht eine Ebene mit zufällig verteilten ionisierten Donatoren befindet, deren Coulomb-Potentiale sich zu dem Zufallspotential überlagern. Wir weisen darauf hin, dass sich die langreichweitigen Fluktuationen dieses Potentials anders verhalten als die kurzreichweitigen. Die kurzreichweitigen klingen mit dem Abstand der Donatorebene von der Ebene des 2DES exponentiell ab, werden aber (bei B = 0) nur schwach durch das 2DES abgeschirmt. Diese Fluktuationen haben wir durch die endlichen Leitfähigkeiten und die Stoßverbreiterung der Landau-Niveaus berücksichtigt. Die langreichweitigen Fluktuationen, andererseits, sind nur schwach von der Entfernung der Donatorebene abhängig, werden aber stark vom 2DES abgeschirmt. Diese sollte man bei der selbstkonsistenten Abschirmungsrechnung explizit berücksichtigen. Erste Versuche in dieser Richtung zeigen, dass sie die Quanten-Hall-Plateaus verbreitern, verschieben und stabilisieren können. Sie sollten besonders bei breiten Proben wichtig werden, bei denen sie zusätzliche inkompressible Streifen im Probeninneren verursachen können. Schließlich diskutieren wir in Kapitel 6 Abschirmungseffekte in einem Doppelschichtsystem aus zwei parallelen 2DES. Interessante neue Effekte treten auf, wenn die Schichten verschiedene Dichten haben. Das Auftreten inkompressibler Streifen in der einen Schicht kann dann drastische Auswirkungen auf die andere Schicht haben. Widerstandsmessungen in Abhängigkeit vom Magnetfeld, die kürzlich an solchen Systemen durchgeführt wurden, zeigen, dass am Rande eines QH-Plateaus Hysterese auftritt, d.h. dass die für ansteigendes Magnetfeld gemessene Kurve nicht mit der für abfallendes Magnetfeld gemessenen Kurve übereinstimmt, wenn dieser Magnetfeldbereich in ein QH-Plateau der anderen Schicht fällt. Wir entwickeln ein Modell und beschreiben Modellrechnungen, die dieses Phänomen plausibel machen.
Cloning and functional characterization of novel genes expressed preferentially in the human retina
(2005)
The human retina is a multi-layered neuronal tissue specialized for the reception and processing of visual information. The retina is composed of a great diversity of neuronal cell types including rod and cone photoreceptors, bipolar cells, ganglion cells, amacrine cells, horizontal cells and Müller glia. In response to light, a coordinated series of molecular events, the so-called phototransduction cascade, is triggered in photoreceptor cells and the signals from the photoreceptors are further processed by the bipolar and ganglion cells to the higher centers of the brain. The retina as highly complex system may be greatly susceptible to genetic defects which can lead to a wide range of disease phenotypes. Therefore, isolation and characterisation of the genes active in the human retina will facilitate our deeper understanding of retinal physiology and mechanisms underlying retinal degeneration and provide novel candidates for the retinal disease genes. To identify novel genes that are specifically or predominantly expressed in the human retina, a cDNA library enriched for retina specific transcripts was generated using suppression subtractive hybridization (SSH) technique. In total, 1113 clones were randomly isolated from the retina SSH cDNA library and partially sequenced. On the basis of BLASTN algorithm analysis these clones were classified into four categories including those with I) significant homology to known human genes (766/1113), II) significant homology to partial transcripts and hypothetical gene predictions (162/1113), III) no homology to known mRNAs (149/1113), and IV) vector sequences and clones derived from mitochondrial genes (36/1113). After correcting for redundancy, category I represented 234 known human genes and category II a total of 92unknown transcripts. Clones from category I, were selected for expression analysis by RT-PCR in a great number of human tissues. This resulted in the identification of 16 genes which were expressed exclusively in the retina, 13 which were highly expressed in the retina compared to other tissues, 12 genes which were specifically expressed in neuronal tissues and 48 ubiquitously expressed genes. Thus, our expression analysis resulted in the identification of 29 genes exclusively or abundantly transcribed in the human retina. Of those, retina specific genes L25,L33, L35, L37, L38 and L40 were selected for further analysis. To characterize the complete mRNA sequences of these transcripts a full-length human retina cDNA library was constructed. The analysis of the L25 gene revealed three splicing variants of the ABCC5 gene, consequently named ABCC5_SV1 (SV1), ABCC5_SV2 (SV2) and ABCC5_SV3 (SV3).These isoforms comprise the first five exons of ABCC5 and additional novel exons named 5a, 5b and 5c, generated by differential exon usage. The determined lengths of the three transcripts are 2039 bp, 1962 bp, and 1887 bp in size, respectively. RT-PCR, real-time PCR and Northern blot analysis of ABCC5 as well as the isoforms SV1, SV2 and SV3demonstrated high levels of expression for all transcripts in the retina compared to other tissues. Analysis of their nucleotide sequences revealed that inclusion of exon 5a in splicing variant SV1 produced a frame shift and premature termination codon (PTC). Our data show that this splice variant is the target of nonsense mediated mRNA decay (NMD). This was shown by inhibition of protein synthesis with antibiotics puromycin and anisomycin in human cell lines A-RPE 19 and Y79. Our analysis resulted in an increase of the PTC containing transcript and a decrease of the ABCC5 transcript. Conversely, the amount of both transcripts (SV1 and ABCC5) returned to pre-treatment levels after removal of the inhibitors. Together, our results suggest that alternative splicing of the ubiquitously expressed ABCC5 gene in addition to NMD is involved in retina-specific transcriptional regulation of the mRNA level of ABCC5. In contrast, additional experiments demonstrated that the levels of expression ofSV2 and SV3 isoforms do not appear to influence ABCC5 transcription. Several of the cloned genes were selected for additional genotyping of single nucleotide polymorphisms (SNPs) in order to construct their SNP maps which are going to be used for future association studies of complex disease AMD. Thus, identification of novel retinal genes and their functional characterization will further our elucidation of retinal physiology in general and in the diseased state in particular, by providing candidate retinal disease genes.