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The main subject of the thesis is the investigation of low-temperature-grown (LTG) GaAs-based photoconductive switches used in the generation of continuous-wave (CW) and pulsed terahertz (THz) radiation. The use of photoconductive switches based on low-temperature-grown GaAs proved to be a viable option in generating electromagnetic transients on a subpicosecond time-scale, corresponding to frequencies of ~1012 Hz (between microwave and far-infrared). The most appealing property of LTG-GaAs is the ultra-short carrier lifetime obtained by incorporation of a large number of As defects when GaAs is grown at low temperatures. However, the reason for poor THz emission efficiency (low CW-THz power lrvrls) is still up to this date not fully understood. The various reasons are to be found in both, optoelectronic properties of the active layer (photoconducting material) as well as in the device characteristics. The thesis focuses primarily on the limitation imposed to the performance of the THz emitters by the material of choice for the active layer (LTG-GaAs) and secondarily, on the impact of a particular emitter design on the THz radiation efficiency. In the beginning of the thesis one finds an ample overview on the electrical and optical properties of the LTG-GaAs material. A special chapter deals with the main features of current-voltage and CW-THz emission characteristics measured from a photoconductive antenna employed as photomixer. We observed deviations from the theoretical predictions of photomixing theory which were explained by considering the high-field electrons effects (velocity overshoot and elongation of the carrier trapping time). With the scope to provide a better understanding of the correlation between device and material properties when the LTG-GaAs material is integrated with a planar antenna (photoswitch), a special THz double-pulse technique (THz-pump and -probe) was implemented. The experimental results assisted by modeling of the double-pulse THz data provide a gainful insight into the ultrafast dynamics of the electrical field and photogenerated carriers. The outcome of the double-pulse experiments is the evidence for long-living carriers in the LTG-GaAs-based photoconductive antenna under applied bias, with a deleterious impact upon the emitter performance (especially for the CW case). Additionally, by measuring the THz transients generated by a constant laser pulse with and without a CW laser background illumination, we obtained further evidence of strong field-screening effects. This phenomenon was also attributed to the existence of long-living space-charge effects. For both cases (pulsed as well as CW) we derived the de-screening time constant. The principal conclusion of the present study is that, besides shortcomings imposed by the THz-circuitry, photomixers based on materials with traps (defects) exhibit great “affinity” for space-charge screening effects with cumulative and therefore long-lived deleterious impact upon device’s performance. An alternative would be the usage of a transient-time limited device where the response time is given by the carrier collection time, possibly with only one type of carrier responsible for THz signal generation.
As if to bear out the tenet of this study, the field of black British literature has been transformed enormously over the last ten years or so, while this book was in the making. And for myself, too, this has been a formative process. During this time I’ve been supported, challenged, and encouraged by more colleagues and friends than I can acknowledge here. ...
Constructive waterfalls
(1911)
The excavation of valleys by waterfalls is one of the best known and most effective processes by which rivers cut down the surface of the earth. The influence of waterfalls is usually regarded as solely destructive, and as always helping to lower the land. They undermine and cut backward the rock faces over which they fall : by this recession they excavate deep gorges ; and the existence of these gorges enables the adjacent country to be lowered to the level of the valIey floors. The waterfalls, moreover, empty any lakes they rnay reach in their retreat, while the ravines below the falls may drain the springs and thus desiccate the neighbouring hihlands. Observations in various countries had suggested to me that waterfalls may sometimes be constructive in stead of destructive, and that they may reserse their usual procedure, advancing instead of retreating, filling valleys instead of excavating them, and forrning alluvial plains and lakes instead of destroying them. The best illustrations I have seen of such advancing, constructive waterfalls are on some rivers of Dalmatia and Bosnia, where they occur in various stages of development. ...
Safety concerns associated with the use of viral vectors in gene therapy applications have attracted considerable attention towards the development of nonviral vectors as alternatives for DNA delivery. While nonviral vectors are commonly not associated with safety problems, they are still very inefficient compared to viral vectors, and require significant improvements to approach the efficiency of their viral counterparts. Meanwhile ligands or single-chain antibody fragments that bind to cell surface receptors for increased and/or specific cellular uptake, endosome escape activities, and nuclear localization sequences (NLSs) to enhance transport of plasmid DNA into the nucleus, have become available that can be incorporated into nonviral vectors to improve their efficacy. However, as gene delivery is a multistep process, the challenge is to incorporate multiple of these functional elements into a single nonviral vector system, while retaining their specific activities. A promising method to attach such entities to plasmid DNA is the use of multifunctional fusion proteins that bind to DNA through a DNA-binding domain. In principle, two types of DNA-binding domains/proteins can be used to anchor additional functional domains or peptides to a plasmid, namely sequence-specific DNA-binding domains, described in the first part of this thesis, or those that bind DNA independent of its sequence, exemplified in the second part of this work by a derivative of the human HMGB2 protein. The first fusion protein constructed and analyzed contained the E. coli LexA repressor as a sequence-specific DNA-binding domain. In addition, this DNA-carrier protein, termed TEL, included a bacterial translocation domain as an integrated endosome escape activity, and human TGF-a for specific targeting to the EGF-receptor (EGFR). TEL was expressed in E. coli and purified under both native and denaturing conditions. Purified, denatured TEL was refolded and subsequently shown to bind specifically to EGFR-expressing cells. However, inclusion of TEL in complexes of plasmid DNA and poly-L-lysine (pL) did not lead to increased gene delivery into EGFR-expressing COS-1 cells. Most likely this was due to the absence of DNA-binding activity of the LexA moiety in TEL. In contrast, native TEL was able to interact specifically with DNA. Nevertheless, since this interaction was rather weak, and refolding of denatured TEL had not resulted in functional activity of all of its protein domains, it seemed unlikely that fusion proteins containing LexA would exhibit gene transfer capabilities superior to those of similar DNA-carrier proteins previously constructed in our group. Further work therefore focused on the use of the E2C-Sp1C protein as an alternative sequencespecific DNA-binding domain. This artificial zinc-finger protein was fused to the single-chain antibody fragment scFv(FRP5), directed against the human ErbB2 growth factor receptor. The resulting 5-E2C fusion protein was expressed in E. coli and purified under native and denaturing conditions. Refolded and native 5-E2C were found to bind specifically to ErbB2-expressing cells, indicating that scFv(FRP5) in 5-E2C was functional in both preparations. In contrast, whereas refolded 5-E2C bound DNA only weakly, significant DNA binding was observed for native 5-E2C. In addition, it could not only be shown that the interaction of native 5-E2C with DNA containing its recognition sequence was specific, but also that this protein was able to bind DNA and recombinant ErbB2 simultaneously, demonstrating the functionality of both domains in native 5-E2C. Despite these encouraging results, the inclusion of native 5-E2C in pL- or polyethyleneimine (PEI)-DNA complexes did not lead to an (5-E2C-specific) enhancement of gene transfer efficiency, irrespective of the presence of the endosome-disruptive reagent chloroquine during transfection. In the second part of this thesis an alternative approach for the development of DNA-carrier proteins for nonviral gene delivery is described, based on human HMGB2, a DNA-binding protein without sequence specificity. HMGB2 contains an acidic C-terminus that has been found to decrease the affinity of the protein for DNA. Therefore, this C-terminal tail was deleted, resulting in an HMGB2-variant consisting of amino acids 1-186. HMGB2186, purified under native conditions from E. coli lysates, was able to interact with DNA and bound to the surface of different cell lines. Importantly, after binding to plasmid DNA HMGB2186 mediated gene delivery into COS-7 cells with higher efficiency than pL. In addition, HMGB2186-mediated gene transfer was strongly enhanced in the presence of chloroquine, indicating that the endocytic pathway was involved in cellular uptake. To improve internalization and intracellular routing of HMGB2186 as a DNA-carrier, a derivative containing the TAT47-57 cell-penetrating peptide (CPP), reported to facilitate cell entry independent of endocytosis, was constructed. Since this peptide also contains an NLS, in addition an HGMB2186-variant containing the SV40-NLS was constructed to investigate the effect of a peptide that has only nuclear localizing properties. Interestingly, the resulting TAT-HMGB2186 and SV40-HMGB2186 fusion proteins displayed DNA-binding activities similar to HMGB2186, but mediated gene delivery into different cell lines clearly more efficiently than the parental molecule. Furthermore, the efficacy of both fusion proteins was enhanced markedly in the presence of chloroquine, an indication that endocytosis was involved in the transfection process mediated by these proteins. This suggests that the increased transfection efficiency observed for TAT-HMGB2186 was more likely due to the NLS function present in the TAT47-57 peptide, rather than to its ‘cell penetrating properties’. Finally, the incorporation of functional peptides derived from human proteins into HMGB2186 was investigated. An uncharged CPP originating from Kaposi-FGF, reported to facilitate efficient cellular uptake of fused protein domains in an endocytosis-independent manner, was fused to HMGB2186 together with the SV40-NLS. Interestingly, the resulting KSV40-HMGB2186 fusion protein bound DNA similarly as previously tested DNA-carrier proteins, but did not mediate enhanced transfection compared to HMGB2186. In addition, the importin-b-binding (IBB) domain derived from human importin-a2 was investigated as a component of a DNA-carrier protein. Since the IBB domain can function as an NLS, it was fused to HMGB2186 resulting in the DNA-carrier protein IBBHMGB2186. Although IBB-HMGB2186 bound DNA in a similar manner as the other HMGB2186-derivatives, gene delivery mediated by IBB-HMGB2186 was only as effective as HMGB2186 mediated transfection, suggesting no significant role of the IBB domain. However, addition of chloroquine resulted in a remarkable enhancement of IBB-HMGB2186-mediated gene transfer, which was now more efficient than with any other HMGB2186-variant tested, and not much lower than gene transfer mediated by PEI, one of the most efficient transfection reagents available to date. To enhance nonviral gene delivery even further, the HMGB2186-based DNA-carrier proteins described in this thesis might now serve as building blocks for novel fusion proteins that include additional complementing activities. In this respect it seems particularly promising that, under conditions of effective end some escape, IBB-HMGB2186, which consists entirely of protein domains of human origin, was the most efficient of all proteins tested in this work.
RcsB is a central transcriptional regulator in enteric bacteria involved in exopolysaccharide (EPS) biosynthesis, in cell division, in the expression of osmoregulated genes, and regulates at least 20 other genes and operons. It is a member of a phosphorelay system and signal transfer is mediated by phosphorylation through the RcsC/YojN phosphorelay. RcsB proteins modified with the phosphorylation mimic BeF3- as shown by its conformational changes and DNA binding properties and resulted phosphorylated RcsB derivatives with sufficient stability. Both, the wild type RcsB protein and the mutant RcsBD11A could be modified with BeF3-. Non-phosphorylated RcsB has been shown to bind as a heterodimer with the coinducer RcsA at the conserved RcsAB box in Rcs regulated promoters. In this study, it has been shown that the modification of RcsB by BeF3 - (I) has a negative effect on its homodimerization, (II) abolishes the complex formation of RcsAB with the RcsAB box as shown by the EMSA and SPR technique. All the effects were found to be reversible by increasing the NaF concentration in the assays presumably leading to the formation of the inactive BeF4 2- salt. This hypothesis of RcsB being modified by BeF3- was also supported by other phosphodonors like ATP and acetyl phosphate, both of them showed the same negative effect on DNA binding by RcsAB heterodimer giving evidence that BeF3- could be used as a phosphorylation mimic. In addition, the phosphorylation mimic BeF3- was found to be a better phosphorylating agent than ATP and acetyl phosphate. This is the first evidence that phosphorylation of RcsB might have a negative effect on the activation of RcsAB regulated operons. Autophosphorylation of RcsB proves that it has the ability to take up phosphoryl groups and the mutant protein also become autophosphorylated with less efficiency or stability than the wild type protein. RcsB probably takes up phosphoryl groups through RcsC -> YojN -> RcsB phosphorelay pathway. To study the interaction among the proteins in this pathway, fluorescence spectroscopy, NMR spectroscopy, and an in vivo ß galactosidase assay were performed by using two domains of RcsC (T-RcsC and R-RcsC), HPt domain of the protein YojN, and RcsB. The interactions between R-RcsC/YojN-HPt and YojN-HPt/RcsB supports the proposed pathway of phosphorylating RcsB. RcsB might also be phosphorylated by YojN-HPt that is phosphorylated by other sensor kinase other than RcsC in a cross-talk mechanism. The phosphorylation of RcsB by YojN-HPt probably has the same negative effect on cps induction as obtained with BeF3 - effect on DNA binding by RcsAB heterodimer.
The biomarker record in two different lakes in central Europe, Lake Albano and Lake Constance, is used to reflect environmental changes and lake system response during the Late Glacial and Holocene. Extractable organic compounds in lake sediments, which can be assigned to their biological source (biomarkers) function as fingerprints of past aquatic or land plant organisms. Using gas chromatography coupled with mass spectrometry, 21 different biomarkers (predominantly steroids and triterpenoids) as well as a variety of n-alkanes, nalkanols, and n-alkanoic acids could be identified in the sediment records of Lake Albano and Lake Constance. In the Holocene sediments of Lake Albano, the distribution of biomarkers such as dinosterol (dinoflagellates), isoarborinol, and diplopterol (aquatic organisms) indicate three biomarker zones: The period between 0-3,800 years BP (zone 3) is characterized by high concentrations of these biomarkers and others such as tetrahymanol and diploptene. Conversely, zone 2 (3,800-6,500 years BP) shows very low concentrations of all autochthonous biomarkers. In zone 1 (6,500–11,480 years BP), dinosterol, isoarborinol, and diplopterol range on a relatively high level, whereas diploptene and tetrahymanol display comparatively low concentrations. The results suggest at least two distinct changes in the predominance of primary producers during the Holocene, which are related to changes in the lake system such as lake mixing and water column stratification. This interpretation is consistent with previous investigations of Lake Albano sediments including pigment and hydrogen index data (Ariztegui et al., 1996b; Guilizzoni et al., 2002). Allochthonous biomarkers such as long-chain n-alkanes, amyrenones and friedelin indicate a development from forest to a more open landscape from 6,000 and 5.000 years BP, respectively. After a period of high concentrations during the first half of the Holocene, all biomarkers derived from deciduous trees exhibit relatively low values until around 1,000 years BP. Again, this is consistent with results from previous pollen investigations (Ariztegui et al., 2000). The sediment core from Upper Lake Constance comprises the Late Glacial and Holocene. It was analysed for biomarkers and inorganic tracers in order to compare the biomarker results with other proxy data from the same core. Magnetic susceptibility (MS) was measured to get a high-resolution stratigraphic framework of the core and to obtain further information about changes of the proportions of allochthonous and autochthonous input. Enhanced concentrations and accumulation rates of dinosterol (biomarker for dinoflagellates) and biogenic calcite give evidence of increasing lake productivity at the beginning of the Holocene followed by a decrease in bioproductivity after around 7,000 years BP. Younger Dryas sediments are characterized by low amounts of both dinosterol and biogenic calcite indicating a low productivity. The comparison of the concentrations and accumulation rates of b-sitosterol and stigmastanol with parameters reflecting lake productivity suggests that both steroids in Lake Constance sediments are mainly derived from terrigenous sources. Biomarkers as well as concentrations and accumulation rates of allochthonous inorganic compounds such as titanium, magnesium and strontium indicate a slightly enhanced allochthonous input after 8,500 years BP. Significant increase of erosive matter input from enhanced soil erosion is not observed before 4,000 years BP. This can be attributed to the combined effects of precipitation increase as a result of climatic deterioration and anthropogenic deforestation which is consistent with observations from other lakes in Central Europe. The MS record of Lake Constance confirms these results by tracing the climatically induced shifts of more intense bioproduction (low MS caused by increased calcite deposition) during the ‘climatic optimum’. This is followed by increasing input of terrigenous sediment compounds during colder and wetter periods which lead to higher MS values in the lake sediments. The occurrence of tetrahymanol in Lake Constance sediments questions the unambiguous use of tetrahymanol as an indicator for water column stratification. Anaerobic organic macroaggregates within the oxygenated, photic zone of the water column have to be considered as a possible living space for anaerobic microorganisms containing tetrahymanol. The direct comparison of two very different lakes Albano and Constance with respect to biomarkers indicating climate or environmental change provides a contribution to the recent biomarker research for a better understanding of biomarkers in lacustrine sediments.