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Institute
- Physikalisches Institut (82)
- Theodor-Boveri-Institut für Biowissenschaften (75)
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- Johns Hopkins School of Medicine (15)
- Johns Hopkins University School of Medicine (5)
- Department of Biomedical Imaging, National Cerebral and Cardiovascular Research Center, Suita, Japan (2)
- Division of Medical Technology and Science, Department of Medical Physics and Engineering, Course of Health Science, Osaka University Graduate School of Medicine, Suita Japan (2)
- Institut for Molecular Biology and CMBI, Department of Genomics, Stem Cell Biology and Regenerative Medicine, Leopold-Franzens-University Innsbruck, Innsbruck, Austria (2)
- International Max Planck Research School Molecular Biology, University of Göttingen, Germany (2)
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ResearcherID
- J-8841-2015 (1)
In 3 Maccabees, kingship as a form of rule is addressed on two levels: On the political level the question about a good king is addressed against the background of Hellenistic understandings of kingship, using the example of Ptolemy IV Philopator. This king is portrayed at the beginning of 3 Maccabees as a successful, positive, Hellenistic ruler, but one whose good rule goes off the rails. This analysis of the ideal of Hellenistic rule (cf. 3 Macc. 3:12-29; 6:24-28; 7:1-9) is then taken to a theological level: the God of Israel is portrayed as the true good king, the Soter who saves his people in their time of greatest trial (6:29, 32; 7:16). By these means the many divine epithets that are a striking feature of 3 Maccabees are incorporated into the narrative (cf. 2:2-3). Thereby 3 Maccabees not only thematises the conflict with a Hellenistic king who exploits his power in diverse ways but also focuses in a concentrated way the notion of a good (Hellenistic) king into the notion of God as king and ruler.
Excitons in the molecular aggregates of chromophores are key participants in important processes such as photosynthesis or the functioning of organic photovoltaic devices. Therefore, the exploration of exciton dynamics is crucial. Here we report on exciton localization during excited-state dynamics of the recently synthesized tetracene trimer [Liu et al., Org. Lett., 2017, 19, 580]. We employ the surface hopping approach to nonadiabatic molecular dynamics in conjunction with the long-range corrected time-dependent density functional tight binding (LC-TDDFTB) method [Humeniuk and Mitrić, Comput. Phys. Commun., 2017, 221, 174]. Utilizing a set of descriptors based on the transition density matrix, we perform comprehensive analysis of exciton dynamics. The obtained results reveal an ultrafast exciton localization to a single tetracene unit of the trimer during excited-state dynamics, along with exciton transfer between units.
Direct Heteroarylation Polymerization (DHAP) is becoming a valuable alternative to classical polymerization methods being used to synthesize π-conjugated polymers for organic electronics applications. In previous work, we showed that theoretical calculations on activation energy (Ea) of the C–H bonds were helpful to rationalize and predict the selectivity of the DHAP. For readers’ convenience, we have gathered in this work all our previous theoretical calculations on Ea and performed new ones. Those theoretical calculations cover now most of the widely utilized electron-rich and electron-poor moieties studied in organic electronics like dithienyl-diketopyrrolopyrrole (DT-DPP) derivatives. Theoretical calculations reported herein show strong modulation of the Ea of C–H bond on DT-DPP when a bromine atom or strong electron withdrawing groups (such as fluorine or nitrile) are added to the thienyl moiety. Based on those theoretical calculations, new cyanated dithienyl-diketopyrrolopyrrole (CNDT-DPP) monomers and copolymers were prepared by DHAP and their electro-optical properties were compared with their non-fluorinated and fluorinated analogues.
Examining the testing effect in university teaching: retrievability and question format matter
(2018)
Review of learned material is crucial for the learning process. One approach that promises to increase the effectiveness of reviewing during learning is to answer questions about the learning content rather than restudying the material (testing effect). This effect is well established in lab experiments. However, existing research in educational contexts has often combined testing with additional didactical measures that hampers the interpretation of testing effects. We aimed to examine the testing effect in its pure form by implementing a minimal intervention design in a university lecture (N = 92). The last 10 min of each lecture session were used for reviewing the lecture content by either answering short-answer questions, multiple-choice questions, or reading summarizing statements about core lecture content. Three unannounced criterial tests measured the retention of learning content at different times (1, 12, and 23 weeks after the last lecture). A positive testing effect emerged for short-answer questions that targeted information that participants could retrieve from memory. This effect was independent of the time of test. The results indicated no testing effect for multiple-choice testing. These results suggest that short-answer testing but not multiple-choice testing may benefit learning in higher education contexts.
This thesis considers a model of a scalar partial differential equation in the presence of a singular source term, modeling the interaction between an inviscid fluid represented by the Burgers equation and an arbitrary, finite amount of particles moving inside the fluid, each one acting as a point-wise drag force with a particle related friction constant.
\begin{align*}
\partial_t u + \partial_x (u^2/2) &= \sum_{i \in N(t)} \lambda_i \Big(h_i'(t)-u(t,h_i(t)\Big)\delta(x-h_i(t))
\end{align*}
The model was introduced for the case of a single particle by Lagoutière, Seguin and Takahashi, is a first step towards a better understanding of interaction between fluids and solids on the level of partial differential equations and has the unique property of considering entropy admissible solutions and the interaction with shockwaves.
The model is extended to an arbitrary, finite number of particles and interactions like merging, splitting and crossing of particle paths are considered.
The theory of entropy admissibility is revisited for the cases of interfaces and discontinuous flux conservation laws, existing results are summarized and compared, and adapted for regions of particle interactions. To this goal, the theory of germs introduced by Andreianov, Karlsen and Risebro is extended to this case of non-conservative interface coupling.
Exact solutions for the Riemann Problem of particles drifting apart are computed and analysis on the behavior of entropy solutions across the particle related interfaces is used to determine physically relevant and consistent behavior for merging and splitting of particles. Well-posedness of entropy solutions to the Cauchy problem is proven, using an explicit construction method, L-infinity bounds, an approximation of the particle paths and compactness arguments to obtain existence of entropy solutions. Uniqueness is shown in the class of weak entropy solutions using almost classical Kruzkov-type analysis and the notion of L1-dissipative germs.
Necessary fundamentals of hyperbolic conservation laws, including weak solutions, shocks and rarefaction waves and the Rankine-Hugoniot condition are briefly recapitulated.
The main focus of this thesis was the processing of different calcium and magnesium phosphate cements together with an optimization of mechanical and biological properties. Therefore, different manufacturing techniques like 3D powder printing and centrifugally casting were employed for the fabrication of reinforced or biomedically improved implants.
One of the main problems during 3D powder printing is the low green strength of many materials, especially when they are only physically bonded and do not undergo a setting reaction. Such materials need post-treatments like sintering to exhibit their full mechanical performance. However, the green bodies have to be removed from the printer requiring a certain stability. With the help of fiber reinforcement, the green strength of printed gypsum samples could be increased by the addition of polymeric and glass fibers within the printing process. The results showed that fiber reinforcement during 3D powder printing is possible and opens up diverse opportunities to enhance the damage tolerance of green bodies as well as directly printed samples. The transfer to biomedically relevant materials like calcium and magnesium phosphate cements and biocompatible fibers would be the next step towards reinforced patient-specific implants.
In a second approach, centrifugally casting derived from construction industries was established for the fabrication of hollow bioceramic cylinders. The aim was the replacement of the diaphysis of long bones, which exhibit a tubular structure with a high density of cortical bone on the fringe. By centrifugation, cement slurries with and without additives could be fabricated to tubes. As a first establishment, the processing parameters regarding the material (e.g. cement composition) as well as the set-up (e.g. rotation times) had to be optimized for each system. In respect of mechanics, such tubes can keep up with 3D powder printed tubes, although the mechanical performance of 3D printed tubes is strongly dependent on printing directions. Additionally, some material compositions like dual setting systems cannot be fabricated by 3D powder printing. Therefore, a transfer of such techniques to centrifugally casting enabled the fabrication of tubular structures with an extremely high damage tolerance due to high deformation ability. A similar effect was achieved by fiber (mesh) addition, as already shown for 3D powder printing. Another possibility of centrifugally casting is the combination of different materials resulting in graded structures to adjust implant degradation or bone formation. This became especially apparent for the incorporation of the antibiotic vancomycin, which is used for the treatment of bacterial implant infections. A long-term release could be achieved by the entrapment of the drug between magnesium phosphate cement layers. Therefore, the release of the drug could be regulated by the degradation of the outer shell, which supports the release into an acidic bacterial environment. The centrifugally casting technique exhibited to be a versatile tool for numerous materials and applications including the fabrication of non-centrosymmetric patient-specific implants for the reconstruction of human long bones.
The third project aimed to manufacture strontium-substituted magnesium phosphate implants with improved biological behavior by 3D powder printing. As the promoting effect of strontium on bone formation and the inhibitory impact on bone resorption is already well investigated, the incorporation of strontium into a degradable magnesium phosphate cement promised a fast integration and replacement of the implant. Porous structures were obtained with a high pore interconnectivity that is favorable for cell invasion and bone ingrowth. Despite the porosity, the mechanical performance was comparable to pure magnesium phosphate cement with a high reliability of the printed samples as quantitatively determined by Weibull statistics. However, the biological testing was impeded by the high degradation rate and the relating ion release. The high release of phosphate ions into surrounding media and the detachment of cement particles from the surface inhibited osteoblast growth and activity. To distinguish those two effects, a direct and indirect cell seeding is always required for degradable materials. Furthermore, the high phosphate release compared to the strontium release has to be managed during degradation such that the adverse effect of phosphate ions does not overwhelm the bone promoting effect of the strontium ions.
The manufacturing techniques presented in this thesis together with the material property improvement offer a diverse tool box for the fabrication of patient-specific implants. This includes not just the individual implant shape but also the application like bone growth promotion, damage tolerance and local drug delivery. Therefore, this can act as the basis for further research on specific medical indications.
Single-molecule fluorescence microscopy in live \(Trypanosoma\) \(brucei\) and model membranes
(2018)
The eukaryotic parasite Trypanosoma brucei has evolved sophisticated strategies to escape
the host immune response and maintain a persistent infection inside a host. One central
feature of the parasite’s defense mechanism relies on the shielding function of their surface
protein coat. This coat is composed of a dense arrangement of one type of glycosylphosphatidylinositol
(GPI)-anchored variant surface glycoproteins (VSGs) which impair the
identification of epitopes of invariant surface proteins by the immune system. In addition
to the importance of understanding the function of the VSG coat and use it as a potential
target to efficiently fight the parasite, it is also crucial to study its biophysical properties as it is not yet understood sufficiently. This is due to the fact that microscopic investigations
on living trypanosomes are limited to a great extent by the intrinsic motility of the parasite.
In the present study, state-of-the-art single-molecule fluorescence microscopy (SMFM)
is introduced as a tool for biophysical investigations in the field of trypanosome research.
The work encompasses studies of VSG dynamics under the defined conditions of an
artificial supported lipid bilayer (SLB). First, the impact of the lateral protein density on
VSG diffusion was systematically studied in SLBs. Ensemble fluorescence after photobleaching
(FRAP) and complementary single-particle tracking experiments revealed that a
molecular crowding threshold (MCT) exists, above which a density dependent decrease
of the diffusion coefficient is measured. A relative quantification of reconstituted VSGs
illustrated that the VSG coat of living trypanosomes operates very close to its MCT and
is optimized for high density while maintaining fluidity. Second, the impact of VSG
N-glycosylation on VSG diffusion was quantitatively investigated. N-glycosylation was
shown to contribute to preserving protein mobility at high protein concentrations. Third,
a detailed analysis of VSG trajectories revealed that two distinct populations of freely
diffusing VSGs were present in a SLB, which is in agreement with the recent finding, that
VSGs are able to adopt two main structurally distinct conformations. The results from
SLBs were further complemented by single-particle tracking experiments of surface VSGs
on living trypanosomes. A high mobility and free diffusion were measured on the cell
surface, illustrating the overall dynamic nature of the VSG coat. It was concluded that
the VSG coat on living trypanosomes is a protective structure that combines density and
mobility, which is supported by the conformational flexibility of VSGs. These features are
elementary for the persistence of a stable infection in the host.
Different hydrogel embedding methods are presented, that facilitated SMFM in immobilized,
living trypanosomes. The hydrogels were found to be highly cytocompatible for one
hour after cross-linking. They exhibited low autofluorescence properties in the spectral
range of the investigations, making them suitable for super-resolution microscopy (SRM).
Exemplary SRM on living trypanosomes illustrated that the hydrogels efficiently immobilized
the cells on the nanometer lever. Furthermore, the plasma membrane organization was studied in living trypanosomes. A statistical analysis of a tracer molecule inside the
inner leaflet of the plasma membrane revealed that specific membrane domains exist, in
which the tracer appeared accumulated or diluted. It was suggested that this distribution
was caused by the interaction with proteins of the underlying cytoskeleton.
In conclusion, SMFM has been successfully introduced as a tool in the field of trypanosome
research. Measurements in model membranes facilitated systematic studies of VSG dynamics
on the single-molecule level. The implementation of hydrogel immobilization
allowed for the study of static structures and dynamic processes with high spatial and
temporal resolution in living, embedded trypanosomes for the first time.
Previous work of our group has established a role of sphingomyelinases in the regulation
of T cell responses to TCR or pathogen stimulation, and this became particularly
evident at the level of actin cytoskeletal dynamics. The formation of lipid membrane
microdomains is crucial for receptor clustering and signal induction, and therefore,
ceramide accumulation by membrane sphingomyelin breakdown is needed for signalling-
complex-assembly. Pathogen-induced overshooting of SMase activation substantially
impacted the formation of membrane protrusions, with T cell spreading as well as
a front/rear polarisation upon CD3/CD28 co-stimulation [103]. On the other hand, NSM
activation is part of the physiological TCR signal [67], indicating that a spatiotemporally
balanced NSM activation is crucial for its physiological function. It involves actin cytoskeletal
reorganisation and T cell polarisation. These two functions are also of central
importance in directional T cell migration and motility in tissues.
This thesis aims on defining the role of NSM in compartmentalisation of the T cell
membrane in polarisation and migration. Therefore, functional studies on the impact of
NSM activity in these processes had to be complemented by the development of tools
to study ceramide compartmentalisation in living T cells.
Involvement of neuronal nitric oxide synthase (NOS-I) PDZ interactions in neuropsychiatric disorders
(2018)
Neuronal nitric oxide (NO) synthase (NOS-I) and its adaptor protein (NOS1AP) have been repeatedly and consistently associated with neuropsychiatric disorders in several genetic association and linkage studies, as well as functional studies. NOS-I has an extended PDZ domain which enables it to interact with postsynaptic density protein 95 (PSD-95) bringing NOS-I in close proximity to NMDA receptors. This interaction allows NMDA receptor activity dependent calcium-influx to activate NOS-I, linking NO synthesis to regulation of glutamatergic signaling pathways. NOS1AP is a PDZ-domain ligand of NOS-I and has been proposed to compete with PSD-95 for NOS-I interaction. Studies performed on post-mortem brain tissues have shown increased expression of NOS1AP in patients with schizophrenia and bipolar disorder, suggesting that increased NOS-I/NOS1AP interactions might be involved in neuropsychiatric disorders possibly through disruption of NOS-I PDZ interactions. Therefore, I have investigated the involvement of NOS-I in different endophenotypes of neuropsychiatric disorders by targeting its specific PDZ interactions in vitro and in vivo. To this end, I used recombinant adeno-associated virus (rAAV) vectors expressing NOS1AP isoforms/domains (NOS1AP-L: full length NOS1AP; NOS1AP-LC20: the last 20 amino acids of NOS1AP-L, containing the PDZ interaction motif suggested to stabilize interaction with NOS-I; NOS1AP-LΔC20: NOS1AP-L lacking the last 20 amino acids; NOS1AP-S: the short isoform of NOS1AP), residues 396-503 of NOS1AP-L (NOS1AP396-503) encoding the full NOS-I interaction domain, and N-terminal 133 amino acids of NOS-I (NOS-I1-133) encoding for the extended PDZ-domain.
Neuropsychiatric disorders involve morphological brain changes including altered dendritic
development and spine plasticity. Hence, I have examined dendritic morphology in primary cultured hippocampal and cortical neurons upon overexpression of constructed rAAV vectors. Sholl analysis revealed that overexpression of NOS1AP-L and NOS1AP-LΔC20 mildly reduced dendritic length/branching. Moreover, overexpression of all NOS1AP isoforms/domains resulted in highly altered spine plasticity including significant reduction in the number of mature spines and increased growth of filopodia. These findings suggest that NOS1AP affects dendritic growth
and development of dendritic spines, which may involve both, increased NOS-I/NOS1AP interaction as well as interaction of NOS1AP with proteins other than NOS-I. Interestingly, the observed alterations in dendritic morphology were reminiscent of those observed in post-mortem brains of patients with neuropsychiatric disorders.
Given the dendritic alterations in vitro, I have examined, whether disruption of NOS-I PDZ interaction would also result in behavioral deficits associated with neuropsychiatric disorders. To this end, rAAV vectors expressing NOS1AP-L, NOS1AP396-503, NOS-I1-133, and mCherry were stereotaxically delivered to the dorsal hippocampus of 6-week-old male C57Bl/6J mice. One week after surgery, mice were randomly separated into two groups. One of those groups underwent three weeks of chronic mild stress (CMS). Afterwards all mice were subjected to a comprehensive behavioral analysis. The findings revealed that overexpression of the constructs did not result in phenotypes related to anxiety or depression, though CMS had an anxiolytic effect independent of the injected construct. Mice overexpressing NOS-I1-133, previously shown to disrupt NOS-I/PSD-95 interaction, showed impaired spatial memory, sensorimotor gating, social interaction, and increased locomotor activity. NOS1AP overexpressing mice showed mild impairments in sensorimotor gating and spatial working memory and severely impaired social interaction. NOS1AP396-503 overexpressing mice also showed impaired social interaction but enhanced sensorimotor gating and reduced locomotor activity. Taken together, these behavioral findings indicate an involvement of NOS-I PDZ interactions in phenotypes associated with positive symptoms and cognitive deficits of psychotic disorders.
In summary, this study revealed an important contribution of NOS-I protein interactions in the development of endophenotypic traits of neuropsychiatric disorders, in particular schizophrenia,
at morphological and behavioral levels. These findings might eventually aid to a better
understanding of NOS-I-dependent psychopathogenesis, and to develop pharmacologically relevant treatment strategies.
Melanoma and Merkel cell carcinoma (MCC) are highly aggressive cancers of the skin that frequently escape immune recognition and acquire resistance to chemotherapeutic agents, which poses a major obstacle to successful cancer treatment. Recently, a new class of therapeutics targeting the programmed cell death-1 (PD-1) immune checkpoint receptor has shown remarkable efficacy in the treatment of both cancers. Blockade of PD-1 on T cells activates cancer-specific immune responses that can mediate tumor regression. The data presented in this Ph.D. thesis demonstrates that PD-1 is also expressed by subsets of cancer cells in melanoma and MCC. Moreover, this work identifies PD-1 as a novel tumor cell-intrinsic growth receptor, even in the absence of T cell immunity. PD-1 is expressed by tumorigenic cell subsets in melanoma patient samples and established human and murine cell lines that also co-express ABCB5, a marker of immunoregulatory tumor- initiating cells in melanoma. Consistently, melanoma-expressed PD-1 downmodulates T effector cell functions and increases the intratumoral frequency of tolerogenic myeloid- derived suppressor cells. PD-1 inhibition on melanoma cells by RNA interference, blocking antibodies, or mutagenesis of melanoma-PD-1 signaling motifs suppresses tumor growth in immunocompetent, immunocompromised, and PD-1-deficient tumor graft recipient mice. Conversely, melanoma-specific PD-1 overexpression enhances tumorigenicity, including in mice lacking adaptive immunity. Engagement of melanoma- PD-1 by its ligand PD-L1 promotes tumor growth, whereas melanoma-PD-L1 inhibition or knockout of host-PD-L1 attenuates growth of PD-1-positive melanomas. Mechanistically, the melanoma-PD-1 receptor activates mTOR signaling mediators, including ribosomal protein S6. In a proof-of-concept study, tumoral expression of phospho-S6 in pretreatment tumor biopsies correlated with clinical responses to anti-PD-1 therapy in melanoma patients. In MCC, PD-1 is similarly co-expressed by ABCB5+ cancer cell subsets in clinical tumor specimens and established human cell lines. ABCB5 renders MCC cells resistant to the standard-of-care chemotherapeutic agents, carboplatin and etoposide. Antibody-mediated ABCB5 blockade reverses chemotherapy resistance and inhibits tumor xenograft growth by enhancing chemotherapy-induced tumor cell killing. Furthermore, engagement of MCC-expressed PD-1 by its ligands, PD-L1 and PD-L2, promotes proliferation and activates MCC-intrinsic mTOR signaling. Consistently, antibody- mediated PD-1 blockade inhibits MCC tumor xenograft growth and phosphorylation of mTOR effectors in immunocompromised mice. In summary, these findings identify cancer cell-intrinsic functions of the PD-1 pathway in tumorigenesis and suggest that blocking melanoma- and MCC-expressed PD-1 might contribute to the striking clinical efficacy of anti-PD-1 therapy. Additionally, these results establish ABCB5 as a previously unrecognized chemoresistance mechanism in MCC.