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- Üceyler, Nurcan (1)
- Ünzelmann, Maximilian (1)
- Üreyen, Soner (1)
- İşbilir, Ali (1)
Institute
- Graduate School of Life Sciences (1022)
- Theodor-Boveri-Institut für Biowissenschaften (798)
- Physikalisches Institut (412)
- Medizinische Klinik und Poliklinik I (315)
- Institut für Pharmazie und Lebensmittelchemie (294)
- Institut für Anorganische Chemie (261)
- Klinik und Poliklinik für Allgemein-, Viszeral-, Gefäß- und Kinderchirurgie (Chirurgische Klinik I) (258)
- Klinik und Poliklinik für Psychiatrie, Psychosomatik und Psychotherapie (234)
- Institut für Organische Chemie (224)
- Medizinische Klinik und Poliklinik II (218)
Schriftenreihe
Sonstige beteiligte Institutionen
- Fraunhofer-Institut für Silicatforschung ISC (8)
- Helmholtz Institute for RNA-based Infection Research (HIRI) (7)
- Universitätsklinikum Würzburg (5)
- Fraunhofer Institut für Silicatforschung ISC (3)
- Fraunhofer-Institut für Silicatforschung (3)
- Klinikum Fulda (3)
- König-Ludwig-Haus Würzburg (3)
- Technische Hochschule Nürnberg Georg Simon Ohm (3)
- Universitätsklinikum Münster (3)
- CHC Würzburg (Comprehensive Hearing Center) (2)
ResearcherID
- B-1911-2015 (1)
- B-4606-2017 (1)
- C-2593-2016 (1)
- D-1250-2010 (1)
- D-3057-2014 (1)
- I-5818-2014 (1)
- J-8841-2015 (1)
- M-1240-2017 (1)
- N-2030-2015 (1)
- N-3741-2015 (1)
EU-Project number / Contract (GA) number
- 311781 (1)
- 320377 (1)
- EU (FP7/ 2007-2013) (1)
Diese retrospektive, naturalistische Studie beschäftigte sich mit dem Therapeutischen Drug Monitoring von Kindern und Jugendlichen unter Psychopharmakotherapie mit Escitalopram und Mirtazapin. Die Datenauswertung erfolgte anhand von klinischen Routinedaten aus dem TDM-Service des Speziallabors für TDM des Zentrums für psychische Gesundheit des Universitätsklinikums Würzburg. In der Studie wurden die Zusammenhänge zwischen Dosis, Serumkonzentration und positiver bzw. negativer klinischer Effekte, auch im Hinblick auf mögliche Einflussfaktoren wie Geschlecht, Alter, BMI-Status, Komedikation und Raucherstatus, untersucht. Ein weiteres Ziel der Arbeit war, Hinweise für die Definition eines altersspezifischen therapeutischen Referenzbereiches (Diagnoseübergreifend und Depressions-spezifisch) für Kinder und Jugendliche beider Medikamente zu gewinnen. Hierfür wurden für Escitalopram 41 Patienten im Alter zwischen elf und 18 Jahren und für Mirtazapin 23 Patienten im Alter von sechs bis 18 Jahren eingeschlossen und Daten zur Demographie, Serumkonzentrationsbestimmungen im Steady State, Schwere der Erkrankung (CGI-S), Therapieeffektivität (CGI-I) und Nebenwirkungen (UKU-Skala) ausgewertet.
Escitalopram: Die mittlere Tagesdosis betrug 14,8 mg, wobei die Serumkonzentrationen mit einer mittleren Konzentration von 32,2 ng/ml (SD= 26,6 ng/ml) zwischen sechs und 109 ng/ml schwankten. Bei 63,4 % der Patienten lag die ermittelte Serumkonzentration in dem für Erwachsene definierten therapeutischen Referenzbereich (15-80 ng/ml). Zwischen der Tagesdosis und der Serumkonzentration ergab sich eine auf dem 1 %-Niveau signifikante positive lineare Beziehung (rs= 0,46; p= 0,003). 65,9 % der Patienten respondierten seit Behandlungsbeginn. Zwischen der Serumkonzentration und dem therapeutischen Effekt (rs= 0,193; p= 0,282) und zwischen der Serumkonzentration und den Nebenwirkungen (r= 0,127; p= 0,467) konnte jeweils kein signifikanter Zusammenhang gefunden werden. Die Nebenwirkungsrate lag bei 25,7 %, wobei am häufigsten Spannung und innere Unruhe dokumentiert wurde. Mit der Idee, die Definition für den vorläufigen therapeutischen Referenzbereich sowohl der Konsensus-Leitlinie der AGNP (Hiemke et al., 2018) als auch von Hiemke (2019) zu berücksichtigen, wird als vorläufiger therapeutischer Referenzbereich für Escitalopram für Kinder und Jugendliche mit Depression eine untere Grenze zwischen 10 ng/ml bis 15 ng/ml und eine obere Grenze von 50 ng/ml vorgeschlagen. Dieser Bereich liegt niedriger als der für erwachsene Patienten definierte Bereich für Escitalopram von 15 bis 80 ng/ml.
Mirtazapin: Die mittlere Tagesdosis betrug 28,6 mg, wobei die Serumkonzentrationen mit einer mittleren Konzentration von 40,8 ng/ml (SD= 28,1 ng/ml) zwischen 13 und 130 ng/ml schwankten. Für 52,2 % der Patienten lag die Serumkonzentration in dem für Erwachsene definierten therapeutischen Referenzbereich (30-80 ng/ml). Zwischen der Tagesdosis und der Serumkonzentration ergab sich eine auf dem 1 %-Niveau signifikante positive Korrelation (rs= 0,655; p= 0,001). Hinsichtlich des Therapieeffektes respondierten 52,2 % der Patienten seit Behandlungsbeginn. Zwischen der Serumkonzentration und dem therapeutischen Effekt ergab sich ein auf dem 5 %-Niveau signifikanter positiver Zusammenhang (rs= 0,534; p= 0,015). Zwischen der Serumkonzentration und den Nebenwirkungen konnte kein signifikanter Zusammenhang gefunden werden (r= 0,240; p= 0,282). Die Nebenwirkungsrate lag bei 30,4 %, wobei Schläfrigkeit und Sedierung am häufigsten berichtet wurden. Als vorläufiger therapeutischer Referenzbereich für Mirtazapin für Kinder und Jugendliche mit Depression wird eine untere Grenze zwischen 15 ng/ml bis 20 ng/ml und eine obere Grenze von 50 ng/ml vorgeschlagen. Dieser Bereich liegt niedriger als der für erwachsene Patienten definierte Bereich für Mirtazapin von 30 bis 80 ng/ml.
Die Limitationen der vorliegenden naturalistischen Studie beachtend, sollten die Ergebnisse mit Vorsicht interpretiert und anhand einer größeren Stichprobe unter kontrollierteren Bedingungen überprüft werden.
p97 uses the energy of ATP hydrolysis to unfold and thereby segregate proteins. It is involved in various cellular processes such as proteasomal degradation, DNA damage repair, autophagy, and endo-lysosomal trafficking. The specificity for these processes is controlled by more than 30 regulatory cofactors.
Interactions of p97 with cofactors and target proteins are known to be highly dynamic and transient. To identify new interaction partners and to uncover novel cellular functions of p97, the interactome of endogenous p97 was determined by using in cellulo crosslinking followed by immunoprecipitation and mass spectrometry. Myoferlin (MYOF) was identified as a novel interactor of p97 and the interaction was validated in reciprocal immunoprecipitation experiments for different cell lines.
The ferlin family member MYOF is a tail-anchored membrane protein containing multiple C2 domains. MYOF is involved in various membrane repair and trafficking processes such as the endocytic recycling of cell surface receptors. The MYOF interactome was determined by mass spectrometry. Among others, the p97 cofactor PLAA, CD71 and Rab14 were identified as common interactors of p97 and MYOF. Immunoprecipitation experiments with PLAA KO cells revealed that the interaction between MYOF and p97 depends on PLAA. Immunofluorescence microscopy showed a co-localization of MYOF with Rab14 and Rab11, which are both involved in endocytic recycling pathways. Furthermore, immunofluoroscence experiments revealed that MYOF and the p97 cofactor PLAA are localized to Rab14- and Rab5-positive endosomal compartments.
Using p97 inhibitors and p97 trapping mutants, the presence of p97 at MYOF-positive and Rab14-positive structures could be demonstrated. Consistent with this finding, the endocytic recycling of transferrin was delayed upon inhibition of p97. Taken together, this work identified MYOF as a novel interactor of p97 and suggests a role for p97 in the recycling of endocytic cargo.
Trotz zahlreicher medizinischer Fortschritte sind Krebserkrankungen weiterhin eine der häufigsten Todesursachen weltweit (Bhupender S. Chhikara und Keykavous Parang 2023). Dabei ist das Multiple Myelom (MM) die zweithäufigste Krebserkrankung des blutbildenden Systems und für ca. 20% aller Todesfälle durch hämatologische Erkrankungen verantwortlich (Derlin und Bannas 2014). Die Krankheit gilt als schwer heilbar, die Patienten leiden unter schwerwiegenden Symptomen und die aktuelle Standardtherapie mittels hochdosierter Chemotherapeutika geht mit starken Nebenwirkungen einher (Cowan et al. 2022). Insofern besteht ein großes Interesse daran, neue und ergänzende Behandlungsmethoden zu finden. In der Forschung etablierte und bereits mit vielversprechenden Ergebnissen an Menschen mit anderen Krebserkrankungen getestete Verfahren sind die Methionin- (MetR) (Kaiser 2020) bzw. Cysteinrestriktion (CysR) (Garcia-Bermudez et al. 2020). Deshalb wurde in der vorliegenden Arbeit in vitro überprüft, ob diese Methoden grundsätzlich einen möglichen Ansatz für die Therapie des MM darstellen. Untersucht wurden die murine MPC11- sowie die humanen L363- und KMS12-BM-Zelllinien des MM. Dabei konnte die antiproliferative Wirkung der Restriktionen bestätigt werden. Darüber hinaus wurde nachgewiesen, dass der Mangel der Aminosäuren nicht über endogene Stoffwechselwege kompensiert werden kann, die Zellen also von der exogenen Zufuhr abhängig sind. Des Weiteren wurde das Metabolom der MPC11-Zellen unter MetR massenspektrometrisch analysiert und ein charakteristischer „metaboler Fingerabdruck“ erstellt. Die Proliferationshemmung ohne Rückgang der Zellzahlen unter den Anfangswert zusammen mit den die gesunde Morphologie der Zellen dokumentierenden EVOS Bildern belegt das Vorliegen eines Low-Energy-Metabolismus (LEM) ohne Absterben der Zellen (Schmitz et al. 2021a). Als geeignete Marker (charakteristischer „metabolen Fingerabdruck“) für einen MetR induzierten LEM eignen sich das bereits in L929-Zellen herauskristallisierte Absinken des Kreatins, der Aminosäuren und Purine bzw. Pyrimidine sowie der Anstieg des Acetoacetats. In den MPC11-Zellen kommen zusätzlich eine Zunahme der Folsäure und eine Abnahme des SAM, SAH und MTA aus dem Methionin- bzw. MTA-Zyklus, des Pentose-5-Phosphats aus dem Pentose-Phosphat-Weg und des Hexose- und Glyceralphosphats sowie des Pyruvats und des Laktats aus der Glykolyse hinzu. Kein klassischer Marker für einen LEM, aber aufgrund des signifikanten Anstiegs dennoch als auffällige Abweichung unter MetR zusätzlich erwähnenswert, ist der deutliche Anstieg des Cystins.
The Macroeconomic Dimensions of Credit: A Comprehensive Analysis of Finance, Inequality and Growth
(2024)
Schumpeter's monetary growth theory is particularly influential for the modern understanding of the macroeconomic role of banks and credit. Based on this theory, this dissertation examines the macroeconomic role of the financial system, especially credit, in (1) generating economic growth, (2) directing economic resources and (3) distributing wealth.
Chapter 3 first shows empirically that 1) there is a positive correlation between the growth of credit and economic growth, even for developed countries, 2) no empirical correlation between household saving and economic growth can be established, and 3) there are both positive, negative and insignificant effects of credit on economic growth at country-specific level. Thus, there is broad empirical support for Schumpeter's monetary hypotheses.
A particularly interesting application of Schumpeter's growth theory can be seen in China. The results of the empirical study suggest that there is generally a positive correlation between credit and economic growth in China, that is, however, not linear in terms of regions, time and size of the financial system. Furthermore, the results in Chapter 4 suggest that credit-financed industrial policy in China may have contributed to more investment and GDP growth, although there are non-linearities between individual industries and types of companies.
Finally, Chapter 5 raises the question of the role of the financial system in the distribution of wealth. While credit to households and companies, together with indicators of working and saving behavior and the age structure of the population, are the most important determinants of wealth inequality, there are also various non-linearities in the relationship between credit and wealth inequality, including in relation to the level of development of financial systems and home ownership ratios.
Binge Eating Disorder (BED) is a common, early-onset mental health condition characterised by uncontrollable episodes of overeating followed by negative emotions such as guilt and shame. An improved understanding of the neurocognitive mechanisms underlying BED is central to the development of more targeted and effective treatments. This thesis comprises a systematic review and three empirical studies contributing to this endeavour.
BED can be thought of as a disorder of cognitive-behavioural control. Indeed, self-report evidence points towards enhanced impulsivity and compulsivity in BED. However, retrospective self-reports do not capture the mechanisms underlying impulsive and compulsive lapses of control in the moment. The systematic review therefore focussed on the experimental literature on impulsivity and compulsivity in BED. The evidence was very mixed, although there was some indication of altered goal-directed control and behavioural flexibility in BED. We highlight poor reliability of experimental paradigms and the failure to properly account for weight status as potential reasons for inconsistencies between studies. Moreover, we propose that impulsivity and/or compulsivity may be selectively enhanced in negative mood states in BED and may therefore not be consistently detected in lab-based studies.
In the empirical studies, we explored the role of behavioural flexibility in BED using experimental and neuroimaging methods in concert with computational modelling. In the first empirical study, we assessed the reliability of a common measure of behavioural flexibility, the Probabilistic Reversal Learning Task (PRLT). We demonstrate that the behavioural and computational metrics of the PRLT have sufficient reliability to justify past and future applications if calculated using hierarchical modelling. This substantially improves reliability by reducing error variance. The results support the use of the PRLT in the second and third empirical studies on development and BED.
Because a majority of patients develop BED as adolescents or young adults, we speculated that it may emerge as a consequence of disrupted or deficient
maturation of behavioural flexibility. Little is known about typical development in this domain. We therefore investigated normative development of reversal learning from adolescence to adulthood in the second empirical study. Typically- developing adolescents exhibited less adaptive and more erratic and explorative behaviour than adults. This behaviour was accounted for by reduced sensitivity to positive feedback in a reinforcement learning model, and partially mediated by reduced activation reflecting uncertainty in the medial prefrontal cortex, a region known to mature substantially during adolescence.
In the third empirical study, we investigated reversal learning in BED, paying special attention to potential biases associated with learning from wins vs learning from losses. We speculated that negative urgency could make it more difficult for BED patients to learn and make decisions under pressure to avoid losses. To dissociate between effects of excess weight and BED, we collected data from obese individuals with and without BED as well as normal-weight controls. As hypothesised, there were subtle neurocognitive differences between obese participants with and without BED with regard to learning to obtain rewards and to avoid losses. Obese individuals showed relatively impaired learning to obtain rewards, while BED patients showed relatively impaired learning to avoid losses. This was reflected in differential learning signals in the brain and associated with BED symptom severity.
In sum, this thesis shows that the evidence on impulsivity and compulsivity in BED is inconsistent and offers potential explanations for this inconsistency. It highlights the need for reliability in interindividual difference research and indicates ways to improve it. Further, it charts the typical development of reversal learning from adolescence to adulthood and underscores the relevance of exploration in the context of learning and decision-making in adolescence. Finally, it demonstrates qualitative differences between BED and obesity, hinting at a pivotal role of aversive states in loss of control in BED.
Allogeneic hematopoietic cell transplantation (Allo-HCT) is the main and only treatment for many malignant and non-malignant haematological disorders. Even though the treatment has improved through the years and patient life expectancy has increased, graft versus host disease (GvHD) is still considered the main obstacle and one of the main reasons for increased mortality. Furthermore, improved patient’s survival and life expectancy brought into question the late post-HCT complications. The leading cause of late death after allo-HCT is the relapse of primary disease as well as chronic GvHD (cGvHD). However, a clear relationship was also described with pulmonary complications, endocrine dysfunction and infertility, and cataracts in post-HCT patients. In the last years big concern regarding a cumulative cardiovascular incidence in long-term survivors has been raised. Severe cardiovascular disease (CVD) is caused by atherosclerosis which is considered a chronic inflammatory disease of blood vessels. As such, it takes a long time from endothelial damage, as the onset event, and followed plaque formation to a manifestation of severe consequences, such as stroke, coronary heart disease, or peripheral arterial disease. Endothelial damage is well documented in patients post-HCT. In the context of allo-HCT, the endothelial damage is induced by the conditioning regimen with or without total body irradiation (TBI). Furthermore, endothelial cells (ECs) have been documented as a target of GvHD and increased concentration of circulating endothelial cells (CEC) coinciding with an increase in the number of circulating alloreactive T cells. According to 2021 ESC Guidelines on CVD prevention, the main atherosclerotic CVD (ASCVD) risk factors are blood apolipoprotein B (ApoB)-containing lipoproteins (of which low-density lipoprotein (LDL) is the most abundant), high blood pressure, cigarette smoking and diabetes mellitus (DM). GvHD is considered a high-risk factor for the onset of dyslipidaemia, hypertension, and DM. Overall, the risk of premature cardiovascular death is 2.7 fold increased in comparison to the general population, while the cumulative incidence of cardiovascular complications was shown to be up to 47% at ten years after reduced intensity conditioning (RIC), post-HCT. However, up to date, there are no available studies elucidating the interconnection between GvHD and atherosclerosis. The goal of this study was, therefore, to investigate the involvement of GvHD in the progression of atherosclerosis as well as to elucidate whether cytotoxic, CD8+ T cells that were shown to play a significant role in endothelial damage during the course of skin GvHD on one hand, and inducers of formation of unstable plaque on the other, are involved in this interconnection. For that purpose we established a novel minor histocompatibility anti gens (miHAg) allo-HCT Western diet (WD)-induced atherosclerosis mouse model. We were able to show that GvHD has a significant impact on atherosclerosis development in B6.Ldlr−/− recipient mice even in the absence of overt clinical disease activity. It seems that the impact is at least partly induced by CD8+ T cells, that showed significantly increased infiltration of aortic lesions in mice facing subclinical GvHD. As studies have shown in regular atherosclerotic mouse models as well as in humans, these CD8+ T cells exhibited not only increased expression of genes involved in activation, survival and differentiation to cytotoxic phenotype, but also some genes pointing out their exhaustion, that were absent in CD4+ T cell cluster. When anti-CD8β antibody was applied once per week along with WD feeding for eight weeks, the plaque formation was significantly reduced in aorta and aortic root pointing out the importance of these cells in an alloreactivity induced lesion formation. Furthermore, anti-CD8β treatment led to significantly decreased necrotic core formation followed by overall increase in plaque stability. Strikingly, bone marrow plus T cells (BMT) recipients fed WD showed significantly increased serum cholesterol levels in comparison to bone marrow (BM) (a group lacking alloreactive T cells that induce GvHD). This effect was reversed when anti-CD8β treatment was applied, suggesting, at least partly, an impact of alloreactive CD8+ T cells on cholesterol levels. Expression of genes responsible for lipid metabolism pointed out the tendency of the liver to regulate the increased cholesterol levels, however, the mechanism behind this phenotype still remains to be revealed. On the other hand, the impact of obesity, induced by chronic high-fat diet (HFD) feeding, has been shown to be an independent risk factor for gastrointestinal GvHD. Similarly, in major histocompatibility complex (MHC) disparate allo-HCT mouse model, we have noticed that even short-term WD intake leads to a significant decrease in survival of mice post-HCT. When the concentration of transplanted alloreactive T cells was reduced, the survival was improved, pointing out the involvement of these cells in the pathogenesis. Additionally, bioluminescence imaging (BLI) during initiation and effector phase of acute GvHD (aGvHD) revealed increased infiltration of alloreactive T cells in mice fed WD. Studies in an obesity model, we could confirm the involvement of specifically CD4+ T cells in WD induced impact, as the relative number of these cells was significantly increased in small intestine on day six post-HCT in mice fed WD. This increased intestinal infiltration was preceded by increase in the number of alloreactive T cells expressing intestine homing receptor (α4β7 integrin) in peripheral lymph nodes (LNs). Even though the number of T cells was not changed in the spleen of WD fed mice, the subset of CD4+ and CD8+ T cells that were highly secreting TNFα was increased as well as the expression of genes regulating pro-inflammatory cytokines such as IL-6 and interferon (IFN)γ pointing out significant WD-induced inflammation. Moreover, slight tendency towards increased intestinal permeability and load of translocated luminal bacteria, that we observed, could induce severe endotoxemia and dysregulated systemic immune response that could lead to detrimental induction of cell death. Justifying our speculations, we noted increased levels of transaminases and an increase in lactate dehydrogenase (LDH) levels (pointing out significant tissue damages). However, the exact mechanism behind this detrimental WD impact still remains to be elucidated.
OSM, ein Vertreter der IL-6-Typ-Zytokine, ist nicht nur für entzündliche, sondern auch für metabolische Prozesse von Bedeutung. Vorarbeiten der Arbeitsgruppe GEIER/HERMANNS und Studien von KOMORI et al. legen protektive Eigenschaften des Zytokins nahe, da Mäuse, denen OSMR fehlte, Charakteristika des metabolischen Syndroms aufwiesen. Zur eingehenderen Untersuchung der von OSM vermittelten Wirkung auf den murinen Lipidstoffwechsel wurden zwei für die NAFLD und Atherosklerose anfällige Modelle herangezogen und jeweils in Gegenwart und Abwesenheit des Osmr studiert: Weibliche Apoe-/-(Osmr-/-) und Ldlr-/-(Osmr-/-) Mäuse wurden über einen Zeitraum von zwölf Wochen mit westlicher Diät gefüttert, wöchentlich gewogen, am Ende der Diät geopfert und geerntet. Wildtypische C57Bl/6-Mäuse erfuhren die gleiche Behandlung und dienten als Referenzgruppe. Im Rahmen des Promotionsprojektes wurden Leberfettgehalt, Serumlipidspiegel, Lipoproteinfraktionen und Stuhllipide von Apoe-deletierten Mäusen bestimmt und mit bereits vorhandenen Daten der Ldlr-/-(Osmr-/-) und wildtypischen Mäuse in Beziehung gesetzt. Expressionsanalysen von am Lipidstoffwechsel beteiligten Genen in Darm-, Leber- und Fettgewebe trugen dazu bei, OSM-abhängige Regulationen aufzudecken.
Ldlr-/- Tiere nahmen unter der Diät exzessiv zu, hatten hohe Serumspiegel an Leptin, Gluco-se und Lipiden, eine Lebersteatose und, begleitet von einer Induktion des Vldlr, erhöhte inflammatorische Marker im visceralen Fettgewebe. Der zusätzliche Knockout des Osmr ging mit einer geringeren Vldlr-Expression im Fettgewebe und einer hepatozytären Induktion von Cyp7a1 einher und resultierte in einem metabolisch günstigeren Phänotyp. Apoe-defiziente Tiere unterschieden sich hinsichtlich ihrer Gewichtszunahme nicht von Ldlr-/-Osmr-/- und C57Bl/6-Mäusen. Überraschenderweise zeigten sich im Serum von Apoe-/-Osmr-/- jedoch gegenüber Apoe-/- Mäusen erhöhte Konzentrationen des Gesamt- und VLDL-Cholesterins, der Triglyceride und freien Fettsäuren. Obwohl Lebern der Apoe-/-Osmr-/- Mäuse geringere Ldlr- und Lrp1-mRNA-Spiegel als die der Apoe-/- Mäuse aufwiesen, hatten sie einen höheren hepatischen Cholesteringehalt. Bei gesteigerter Cpt1a-Expression fiel der hepatische Tri-glyceridgehalt Apoe-deletierter Mäuse geringer aus als in Ldlr-/-(Osmr-/-) und wildtypischen Tieren. Unter Umgehung einer Fettgewebsentzündung präsentierten Apoe-defiziente Mäuse Hinweise einer inflammatorischen Leberschädigung, die pathogenetisch am ehesten mit einer gestörten Cholesterinhomöostase in Verbindung zu bringen war.
Abhängig vom genetischen Hintergrund des Mausmodells hatte OSM schützende oder schädliche Effekte auf den Lipidmetabolismus. Die Ergebnisse der vorliegenden Arbeit betonen die entscheidende Bedeutung entzündlicher, von OSM modulierter Prozesse für den Fettstoffwechsel in Leber- und Fettgewebe. Weiterführende Experimente sind nötig, um die den Beobachtungen zugrunde liegenden molekularen Mechanismen zu entschlüsseln.
The evolutionary success of higher plants is largely attributed to their tremendous developmental
plasticity, which allows them to cope with adverse conditions. However, because these adaptations
require investments of resources, they must be tightly regulated to avoid unfavourable trade-offs.
Most of the resources required are macronutrients based on carbon and nitrogen. Limitations in the
availability of these nutrients have major effects on gene expression, metabolism, and overall plant
morphology. These changes are largely mediated by the highly conserved master kinase SNF1-RELATED
PROTEIN KINASE1 (SnRK1), which represses growth and induces catabolic processes. Downstream of
SnRK1, a hub of heterodimerising group C and S1 BASIC LEUCINE ZIPPER (bZIP) transcription factors has
been identified. These bZIPs act as regulators of nutrient homeostasis and are highly expressed in
strong sink tissues, such as flowers or the meristems that initiate lateral growth of both shoots and
roots. However, their potential involvement in controlling developmental responses through their
impact on resource allocation and usage has been largely neglected so far. Therefore, the objective of
this work was to elucidate the impact of particularly S1 bZIPs on gene expression, metabolism, and
plant development.
Due to the high homology and suspected partial redundancy of S1 bZIPs, higher order loss-of-function
mutants were generated using CRISPR-Cas9. The triple mutant bzip2/11/44 showed a variety of robust
morphological changes but maintained an overall growth comparable to wildtype plants. In detail
however, seedlings exhibited a strong reduction in primary root length. In addition, floral transition
was delayed, and siliques and seeds were smaller, indicating a reduced supply of resources to the shoot
and root apices. However, lateral root density and axillary shoot branching were increased, suggesting
an increased ratio of lateral to apical growth in the mutant. The full group S1 knockout
bzip1/2/11/44/53 showed similar phenotypes, albeit far more pronounced and accompanied by
growth retardation. Metabolomic approaches revealed that these architectural changes were
accompanied by reduced sugar levels in distal sink tissues such as flowers and roots. Sugar levels were
also diminished in leaf apoplasts, indicating that long distance transport of sugars by apoplastic phloem
loading was impaired in the mutants. In contrast, an increased sugar supply to the proximal axillary
buds and elevated starch levels in the leaves were measured. In addition, free amino acid levels were
increased in bzip2/11/44 and bzip1/2/11/44/53, especially for the important transport forms
asparagine and glutamine. The increased C and N availability in the proximal tissues could be the cause
of the increased axillary branching in the mutants.
To identify bZIP target genes that might cause the observed shifts in metabolic status, RNAseq
experiments were performed. Strikingly, clade III SUGARS WILL EVENTUALLY BE EXPORTED (SWEET)
8
genes were abundant among the differentially expressed genes. As SWEETs are crucial for sugar export
to the apoplast and long-distance transport through the phloem, their reduced expression is likely to
be the cause of the observed changes in sugar allocation. Similarly, the reduced expression of
GLUTAMINE AMIDOTRANSFERASE 1_2.1 (GAT1_2.1), which exhibits glutaminase activity, could be an
explanation for the abundance of glutamine in the mutants. Additional experiments (ATAC-seq, DAPseq, PTA, q-RT-PCR) supported the direct induction of SWEETs and GAT1_2.1 by S1 bZIPs. To confirm
the involvement of these target genes in the observed S1 bZIP mutant phenotypes, loss-of-function
mutants were obtained, which showed moderately increased axillary branching. At the same time, the
induced overexpression of bZIP11 in axillary meristems had the opposite effect.
Collectively, a model is proposed for the function of S1 bZIPs in regulating sink tissue development. For
efficient long-distance sugar transport, bZIPs may be required to induce the expression of clade III
SWEETs. Thus, reduced SWEET expression in the S1 bZIP mutants would lead to a decrease in apoplastic
sugar loading and a reduced supply to distal sinks such as shoot or root apices. The reduction in longdistance transport could lead to sugar accumulation in the leaves, which would then increasingly be
transported via symplastic routes towards proximal sinks such as axillary branches and lateral roots or
sequestered as starch. The reduced GAT1_2.1 levels lead to an abundance of glutamine, a major
nitrogen transport form. The combined effect on C and N allocation results in increased nutrient
availability in proximal tissues, promoting the formation of lateral plant organs. Alongside emerging
evidence highlighting the power of bZIPs to steer nutrient allocation in other species, a novel but
evolutionary conserved role for S1 bZIPs as regulators of developmental plasticity is proposed, while
the generation of valuable data sets and novel genetic resources will help to gain a deeper
understanding of the molecular mechanisms involved
Forests are essential sources of tangible and intangible benefits, but global climate change associated with recurrent extreme drought episodes severely affects forest productivity due to extensive tree die-back. On that, it appeals to an urgency for large-scale reforestation efforts to mitigate the impact of climate change worldwide; however, there is a lack of understanding of drought-effect on sapling growth and survival mechanisms. It is also challenging to anticipate how long trees can survive and when they succumb to drought. Hence, to ensure success of reforestation programs and sustainable forest productivity, it is essential to identify drought-resistant saplings. For that, profound knowledge of hydraulic characteristics is needed. To achieve this, the study was split into two phases which seek to address (1) how the hydraulic and anatomical traits influence the sapling’s growth rate under drought stress. (2) how plant water potential regulation and physiological traits are linked to species’ water use strategies and their drought tolerance.
The dissertation is assembled of two study campaigns carried out on saplings at the Chair of Botany II, University of Würzburg, Germany. The first study involved three ecologically important temperate broadleaved tree species — saplings of 18-month (Acer pseudoplatanus, Betula pendula, and Sorbus aucuparia) — grown from seeds in contrasting conditions (inside a greenhouse and outside), with the latter being subjected to severe natural heat waves. In the second study, two additional temperate species (Fagus sylvatica and Tilia cordata) were added. The drying-out event was conducted using a randomised blocked design by monitoring plant water status in a climate-controlled chamber and a greenhouse.
In campaign I, I present the result based on analysed data of 82 plants of temperate deciduous species and address the juvenile growth rate trade-off with xylem safety-efficiency. Our results indicate biomass production varies considerably due to the contrasted growing environment. High hydraulic efficiency is necessary for increased biomass production, while safety-efficiency traits are decoupled and species-specific. Furthermore, productivity was linked considerably to xylem safety without revealing a well-defined pattern among species. Moreover, plasticity in traits differed between stressed and non-stressed plants. For example, safety-related characteristics were more static than efficiency-related traits, which had higher intra-specific variation. Moreover, we recorded anatomical and leaf traits adjustments in response to a stress condition, but consistency among species is lacking.
In campaign II, I combined different ways to estimate the degree of isohydry based on water potential regulation and connected the iso-anisohydric spectrum (i.e., hydroscape area, HSA) to hydraulic traits to elucidate actual plant performance during drought. We analysed plant water potential regulation (Ψpd and Ψmd) and stomatal conductance of 28-29 month saplings of five species. I used a linear mixed modelling approach that allowed to control individual variations to describe the water potential regulation and tested different conceptual definitions of isohydricity. The combined methods allowed us to estimate species' relative degree of isohydry. Further, we examined the traits coordination, including hydraulic safety margin, HSM; embolism resistance, P88; turgor loss, Ψtlp; stomata closure, Ps90; capacitance, C; cuticular conductance, gmin, to determine time to hydraulic failure (Thf). Thf is the cumulative effect of time to stomata closure (Tsc) and time after stomatal closure to catastrophic hydraulic failure (Tcrit).
Our results show the species' HSA matches their stomatal stringency, which confirms the relationship between stomatal response and leaf water potential decline. Species that close stomata at lower water potential notably had a larger HSA. Isohydric behaviour was mostly associated with leaf hydraulic traits and poorly to xylem safety traits. Species' degree of isohydry was also unrelated to the species' time to death during drying-out experiments. This supports the notion that isohydry behaviours are linked to water use rather than drought survival strategies. Further, consistent with our assumptions, more isohydric species had larger internal water storage and lost their leaf turgor at less negative water potentials. Counter to our expectations, neither embolism resistance nor the associated hydraulic safety margins were related to metrics of isohydry. Instead, our results indicate traits associated with plant drought response to cluster along two largely independent axes of variation (i.e., stomatal stringency and xylem safety). Furthermore, on the temporal progression of plant drought responses, stomatal closure is critical in coordinating various traits to determine species' hydraulic strategies. Desiccation avoidance strategy was linked to Tsc and coordinated traits response of Ps90, Ψtlp, and HSA, whereas desiccation tolerance was related to Tcrit and traits such as lower P88 value, high HSM, and lower gmin. Notably, the shoot capacitance (C) is crucial in Thf and exhibits dichotomous behaviour linked to both Tsc and Tcrit.
In conclusion, knowledge of growth rate trade-offs with xylem safety-efficiency combined with traits linked to species’ hydraulic strategies along the isohydry could substantially enhance our ability to identify drought-resistant saplings to ensure the success of reforestation programs and predicting sensitivity to drought for achieving sustainable forest ecosystems.
Die Erleichterung von einem körperlichen Schmerzreiz besitzt appetitiven Charakter (Leknes et al., 2008; 2011; Seymour et al., 2005), aktiviert belohnungsassoziierte Hirnstrukturen (Leknes et al., 2011; Leknes & Brock, 2014; Leknes & Tracey, 2008; Navratilova & Porreca, 2014) und fördert durch ihre Konditionierbarkeit als Erleichterungslernen bezeichnete appetitive Lern- und Konditionierungsprozesse (Andreatta et al., 2010, 2012; 2013; 2017; Gerber et al., 2014; Tanimoto et al., 2004; Yarali et al., 2008).
Die vorliegende Arbeit bestätigt das angewandte Versuchsparadigma als valides Modell für Erleichterungslernen im Menschen und zeigt erstmals, dass der appetitive Charakter von Schmerzerleichterung auch in Jugendlichen konditionierbar ist. Erfolgreiches Erleichterungslernen zeigte sich dabei in der untersuchten Stichprobe lediglich auf impliziter, nicht aber auf expliziter, kognitiver Ebene. Dies stützt Thesen und vorherige Forschungsbefunde einer Dualität assoziativen Lernens in ein implizites Lernen, welches vornehmlich subkortikale Strukturen erfordert und ein explizites Lernen, das vorrangig kortikale Strukturen wie den präfrontalen Cortex involviert (Andreatta et al., 2010; Strack & Deutsch, 2004; Williams et al., 2001). Die Beobachtungen einer differenten Furcht- versus Erleichterungs-Extinktion bestärken die Thesen eines diversen neuronalen Hintergrunds dieser beiden Lernformen (Diegelmann et al., 2013; Gerber et al., 2014; Yarali et al., 2009; Yarali & Gerber, 2010). Gleichzeitig werfen die Studienergebnisse die Frage auf, ob und inwiefern im Erleichterungslernen von Jugendlichen Unterschiede zu jenem in Erwachsenen bestehen.
Die Hypothese einer verstärkten Akquisition von Erleichterungslernen bei Jugendlichen mit NSSV im Vergleich zu gesunden Jugendlichen ließ sich in der vorliegenden Studie nicht bestätigen. Somit liefern die Ergebnisse keinen direkten Hinweis darauf, dass ein verstärktes Lernen durch Schmerzerleichterung an der Ätiopathogenese von NSSV beteiligt sein könnte. Die vorliegende Arbeit zeigte vielmehr die Tendenz eines abgeschwächten impliziten Erleichterungslernens bei den Jugendlichen mit NSSV. Die tendenziellen Gruppenunterschiede ließen sich nicht hinreichend durch eine differente aktuelle Stimmungslage oder durch eine unterschiedlich starke Ausprägung aversiver emotionaler Anspannungen oder momentaner Angstaffekte erklären. Innerhalb der Gruppe Jugendlicher mit NSSV zeigte sich auch kein Hinweis darauf, dass der Erfolg von Erleichterungslernen vom Schweregrad des NSSV oder von der aktuellen Einnahme von Antidepressiva abhängig sein könnte. Explorative Analysen ergaben, dass Gruppeneffekte in der vorliegenden Studie womöglich aufgrund einer statistischen Unterschätzung, bedingt durch einen zu geringen Stichprobenumfang, nicht das Signifikanzniveau erreichten und dass Unterschiede im Erleichterungslernen von Jugendlichen mit und ohne NSSV tatsächlich sogar noch stärker ausgeprägt sein könnten. Somit sollte die vorliegende Arbeit als Pilotstudie für zukünftige größer angelegte Studien zu Erleichterungslernen bei NSSV betrachtet werden.
Zukünftige Studien erscheinen insbesondere sinnvoll mit Blick auf die hohe klinische sowie gesellschaftliche Relevanz von NSSV für welches, trotz der hohen Prävalenzen und des deutlich erhöhten Morbiditäts- und Mortalitätsrisikos, zum aktuellen Zeitpunkt noch keine hinreichenden Erklärungsmodelle bestehen. Die Studie bestätigte das Vorliegen eines erhöhten Grades aversiver emotionaler Anspannung in Jugendlichen mit NSSV, welcher zuvor nur an Erwachsenen mit einer BPD untersucht und festgestellt worden war (Niedtfeld et al., 2010; Stiglmayr et al., 2005). Die Abnahme negativer Affekte bei den Jugendlichen mit NSSV im Studienverlauf repliziert die Ergebnisse vorheriger Studien, in denen eine Reduktion selbst-berichteter negativer Affekte durch die Beendigung eines Schmerzreizes beobachtet wurde (Bresin et al., 2010; Bresin & Gordon, 2013). Damit bestärken die Studienergebnisse bestehende Erklärungsmodelle für NSSV, welche eine entscheidende Beteiligung der körperlichen Schmerzen und der Schmerzerleichterung bei der Selbstverletzung an der Affektregulation vermuten. Weiterhin wirft die vorliegende Arbeit die Frage auf, welche Rolle eine veränderte Wahrnehmung von Schmerz und Schmerzerleichterung in der Ätiopathogenese von NSSV einnimmt und wie diese sich auf Lernprozesse auswirkt.
Insgesamt erbrächten weitere Erkenntnisse über den potenziellen Zusammenhang von NSSV und abweichendem Erleichterungslernen ein besseres Verständnis für Mechanismen der Entstehung und Aufrechterhaltung von NSSV und böten zudem möglicherweise Ansätze für neue Therapiemöglichkeiten des Störungsbildes.