570 Biowissenschaften; Biologie
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- Honigbiene (2)
- HopZ1a (2)
- Hybrid speciation (2)
- Hybridisation capture (2)
- Hydrogel (2)
- Hydrogele (2)
- Hypoxie (2)
- IC (2)
- IDP (2)
- Iambic/Trochaic Law (2)
- Illumina amplicon sequencing (2)
- Illuminance (2)
- Immobilisierung (2)
- Immunoassay (2)
- InP (2)
- InPZnS (2)
- India (2)
- Individual-based modeling (2)
- Inflammation (2)
- Influenza A virus (2)
- Insect (2)
- Insulin resistance (2)
- Interdigitated electrodes (2)
- Interoception (2)
- Interspecific interactions (2)
- Introgression (2)
- Intuitive eating (2)
- Invagination (2)
- Ionentransport (2)
- Island biogeography (2)
- Isoflavone (2)
- Isolation (2)
- JUB1 (2)
- Japan (2)
- June 2013 flood (2)
- Jurkat cells (2)
- Just so stories (2)
- Kartoffel (2)
- Kernhülle (2)
- Kettle holes (2)
- Klimaänderung (2)
- Koexpression (2)
- Konnektivität (2)
- Korrelationsanalyse (2)
- Körperfett (2)
- Körperzusammensetzung (2)
- LC–MS/MS (2)
- LEA protein (2)
- LEM-domain protein (2)
- LMA (2)
- LPS (2)
- Lake (2)
- Lamin (2)
- Land use (2)
- Land-use history (2)
- Landscape (2)
- Landscape of fear (2)
- Landschaft der Angst (2)
- Large fragment deletion (2)
- Late Quaternary vegetation (2)
- Leaf senescence (2)
- Leopards (2)
- Limnology (2)
- Lipases (2)
- Lipidomics (2)
- Lysophosphatidylcholine (2)
- Lythrum salicaria (2)
- MADS-domain transcription factor (2)
- MAPK (2)
- MED16 (2)
- MHC (2)
- MSAP (2)
- Manganese (2)
- Markov cluster algorithm (2)
- Martini force-field (2)
- Mate choice (2)
- Maus (2)
- Mechanotransduction (2)
- Medicago truncatula (2)
- Meiosis (2)
- Melampyrum pratense (2)
- Membranproteine (2)
- Mesenchymal stem cells (2)
- Metabolismus (2)
- Metabolit (2)
- Metabolite (2)
- Metaboliten (2)
- Metabolites (2)
- Metabolom (2)
- Metal Metabolism (2)
- Metschnikowia (2)
- Microarrays (2)
- Microbial ecology (2)
- Microbiology (2)
- Microcystin (2)
- Microcystis aeruginosa (2)
- Microsatellite analysis (2)
- Mikroorganismen (2)
- Mikrosatelliten (2)
- Mindfulness (2)
- Mitochondrial DNA (2)
- Mitochondrien (2)
- Mitogenome (2)
- Mitogenomes (2)
- Mitose (2)
- Mitosis (2)
- Mixed mating (2)
- Mixed methods (2)
- Modell (2)
- Molekularbiologie (2)
- Molybdänkofaktor (2)
- Monoclonal antibodies (2)
- Monoclonal antibody (2)
- Monte Carlo method (2)
- Morphology (2)
- Multiplex (2)
- Multiplex mutagenesis (2)
- Muntjac (2)
- Mutation rate (2)
- Mutator locus (2)
- Mykotoxine (2)
- NAB (2)
- NE Germany (2)
- NMR spectroscopy (2)
- NMR structure (2)
- NRPS (2)
- NZO (2)
- Nahrungsnetz (2)
- Namibia (2)
- Nanostruktur (2)
- Network clustering (2)
- Netzwerke (2)
- Neurotoxicity (2)
- Nighttime illumination (2)
- Nitrat (2)
- Nitrogen deposition (2)
- Northern Asia (2)
- Norway (2)
- Nostoc punctiforme (2)
- Nucleus (2)
- NutriAct Family Study (2)
- Nutrition (2)
- O-serotyping (2)
- OMICs tools (2)
- Offspring weight (2)
- Ohrid-Prespa region (2)
- Older adults (2)
- Optogenetik (2)
- Organogenesis (2)
- Orthologous Matrix (OMA) Project (2)
- Outcrossing (2)
- Outcrossing rate (2)
- Outdoor enclosure (2)
- PBPK (2)
- PBS1 (2)
- PC-3 cells (2)
- PEI coating (2)
- PKS (2)
- PUFA (2)
- PXR (2)
- Palaeogenetics (2)
- Paläoökologie (2)
- Parasit (2)
- Parasitoid wasp (2)
- Partial Little Square (2)
- Partnership (2)
- Paternity analysis (2)
- Pathogen (2)
- Pathway design (2)
- Peptid (2)
- Periphyton (2)
- Periplaneta americana (2)
- Personality (2)
- Pex1 (2)
- Pex6 (2)
- Pflanze (2)
- Pflanzengemeinschaften (2)
- Pflanzenwachstum (2)
- Pflanzenökologie (2)
- PhIP (2)
- Phase variation (2)
- Phenole (2)
- Phosphat (2)
- Photosynthesis (2)
- Phylogenetics (2)
- Phylogenie (2)
- Phylogenomics (2)
- Pichia pastoris (2)
- Pinus sylvestris (2)
- Pipistrellus nathusii (2)
- Plankton (2)
- Plant community model (2)
- Plant development (2)
- Plant-soil feedback (2)
- Plants (2)
- Plasma membrane (2)
- Plastid (2)
- Poecilia formosa (2)
- Poecilia latipinna (2)
- Polyadenylierung (2)
- Polyelektrolyt-Multischichten (2)
- Polyethylenglykol (2)
- Polysaccharide (2)
- Population dynamics (2)
- Predator-prey interactions (2)
- Pregnancy (2)
- Priming (2)
- Promiscuous enzymes (2)
- Promotor (2)
- Prostaglandin E2 (2)
- Proteinaggregation (2)
- Proteins (2)
- Pseudomonas aeruginosa (2)
- Puumala virus seroprevalence (2)
- QDs (2)
- QTL mapping (2)
- R Shiny (2)
- RNA (2)
- RNA interference (2)
- RNA virus (2)
- RNA-guided Cas9 (2)
- RT-PCR (2)
- RUNX2 (2)
- Randomized-controlled trial (2)
- Raphidiopsis (2)
- Recombination (2)
- Redox marker (2)
- Redoxreaktion (2)
- Redoxregulation (2)
- Regulation (2)
- Rezeptor (2)
- Rhizophagus irregularis (2)
- Rice (2)
- Riesenvesikel (2)
- Rodent (2)
- Roridula gorgonias (2)
- Rotifera (2)
- RpoS (2)
- RubisCO (2)
- Russia (2)
- S-XRF (2)
- SELEX (2)
- SERS enhancement factor (2)
- SFON (2)
- SIMS techniques (2)
- SPB (2)
- SULT1A1 (2)
- SWL (2)
- Sailfin molly (2)
- Salmonella Typhimurium (2)
- Saprolegnia (2)
- Saprolegniaceae (2)
- Savannas (2)
- Savanne (2)
- Schleudertrauma (2)
- Schmalwand <Arabidopsis> (2)
- Schwefel (2)
- Scopoletin (2)
- Seasonality (2)
- Sediment (2)
- Selenium (2)
- Senescence (2)
- Sequenzierung (2)
- Sequenzierung der nächsten Generation (2)
- Serine cycle (2)
- Sexual selection (2)
- Siberian tree line (2)
- Simulation (2)
- Simulationsmodell (2)
- Smpd1 (2)
- Soil (2)
- Soil function (2)
- Solanaceae (2)
- Solanum lycopersicum (2)
- Space use (2)
- Specific wood density (2)
- Stickstoff (2)
- Strategic growth adjustment (2)
- Störung (2)
- Sulforaphan (2)
- Sulfotransferase (2)
- Sundaland (2)
- Sus scrofa (2)
- Systems Biology (2)
- Säugetiere (2)
- TAS2R (2)
- TEM (2)
- TMS (2)
- TTR (2)
- Tailspike protein (2)
- Tanzania (2)
- Tas2r (2)
- Tas2rs (2)
- Temperate forest (2)
- Tenebrio molitor larvae (2)
- Tocopherol (2)
- Toll and Imd pathways (2)
- Tomate (2)
- Tomato (2)
- Transcription factors (2)
- Transcriptional memory (2)
- Transcriptome assembly (2)
- Transporters (2)
- Tree allometry (2)
- Turkey (2)
- TusA (2)
- Type 2 Diabetes (2)
- UV radiation (2)
- Ulcerative colitis (2)
- Urbanization (2)
- Ursus arctos (2)
- V-ATPase (2)
- Venom proteins (2)
- Verhalten (2)
- Verkhoyansk mountains (2)
- Virus (2)
- Wachstum (2)
- Walker A motif (2)
- Wastewater (2)
- Weakly electric fish (2)
- Weizen (2)
- Weißstorch (2)
- Wildschwein (2)
- Wind (2)
- Wood specific gravity (2)
- Woody aboveground biomass (2)
- X-ray structure (2)
- XMRV (2)
- Xpr1 (2)
- Yap1/Wwtr1 (Taz) (2)
- Zebrafish (2)
- Zelladhäsion (2)
- Zellkern (2)
- Zellweger (2)
- Zellweger syndrome spectrum disorder (ZSSD) (2)
- Zinypr-1 (2)
- Zweizustandsmodell (2)
- Zytoskelett (2)
- aberrations (2)
- accelerometer (2)
- acclimation (2)
- acid mine drainage (2)
- acoustic communication (2)
- actin polymerization (2)
- activated carbon (2)
- activator–inhibitor models (2)
- adaptive introgression (2)
- adaptive processes (2)
- adaptive radiation (2)
- adhesion (2)
- adiponectin (2)
- age (2)
- agricultural landscape (2)
- agrin (2)
- agro-infiltration (2)
- aldehyde oxidoreductase (2)
- alignment (2)
- alignment sensitivity / specificity (2)
- alkylierte polyzyklische aromatische Kohlenwasserstoffe (2)
- alkylphospholipids (2)
- all-cause mortality (2)
- allocation (2)
- alphaherpesvirus (2)
- alternative splicing (2)
- amino acid (2)
- amphibians (2)
- animal cognition (2)
- animal migration (2)
- anthropogenic environment (2)
- anthropogenic food subsidies (2)
- anthropometric measures (2)
- anthropometry (2)
- anti-oxidative response (2)
- antibody producing cell selection (2)
- antioxidant capacity (2)
- anxiety-like behavior (2)
- apis mellifera (2)
- apple (2)
- aquatic ecosystems (2)
- aqueous dispersion (2)
- aqueous-solution (2)
- arable land (2)
- arable weeds (2)
- arbuscular mycorrhizal symbiosis (2)
- archival DNA (2)
- area-based conservation (2)
- arsenic (2)
- arthropod (2)
- artificial light at night (ALAN) (2)
- ascorbate peroxidase (2)
- assembly (2)
- assembly processes (2)
- assembly rules (2)
- autocorrelation (2)
- automated radio telemetry (2)
- axillary bud (2)
- baltic sea (2)
- basal body (2)
- base excision repair (incision activity) (2)
- bat fatalities (2)
- behavior (2)
- behavioral plasticity (2)
- behavioral type (2)
- behavioural syndrome (2)
- benzylic alcohol (2)
- benzylischer Alkohol (2)
- beta diversity (2)
- bias (2)
- bifurcation (2)
- bifurcation theory (2)
- biodiversity conservation (2)
- biodiversity decline (2)
- biodiversity exploratories (2)
- biodiversity facets (2)
- biofortification (2)
- biogene Amine (2)
- biological age (2)
- biological control (2)
- biological invasions (2)
- biological soil crusts (2)
- biomarker detection (2)
- biomimetic sensors (2)
- biosignatures (2)
- biostimulant (2)
- biosynthesis (2)
- biotic filtering (2)
- biotin sulfoxide reductase (2)
- bird migration (2)
- bisphosphonates (2)
- bitter (2)
- bitter taste receptors (2)
- body composition (2)
- body proportions (2)
- bone mineral density (2)
- bone pathologies (2)
- brackish waters (2)
- brain insulin signaling (2)
- branched chain amino acids (2)
- branching (2)
- breeding (2)
- buffer zones (2)
- bush encroachment (2)
- cAMP (2)
- cTBS (2)
- cadmium-free (2)
- caffeine (2)
- calcination (2)
- calcite (2)
- calcium (2)
- calcium carbonate inclusions (2)
- calcium imaging (2)
- calcium influx (2)
- canalization (2)
- cancer epidemiology (2)
- capture enrichment (2)
- carbohydrates (2)
- carbon dots (2)
- carbon isotopes (2)
- carbon labeling (2)
- cardiac development (2)
- cardiomyocyte (2)
- cardiomyogenic differentiation (2)
- carotenoid biosynthesis (2)
- carrion ecology (2)
- cascading effects (2)
- cave fish (2)
- cell adhesion (2)
- cell shape (2)
- cell signaling (2)
- cells (2)
- cellular bioenergetics (2)
- cellular bioimaging (2)
- cellulose polymeric organic matter (2)
- cerami-des (2)
- ceramides (2)
- cereal leaf beetle (2)
- cereal meals (2)
- chemostat experiments (2)
- child growth (2)
- chimeric transcription factors (2)
- chlamydomonas (2)
- chlorophyll a (2)
- chloroplast (2)
- chloroplasts (2)
- chronic diseases (2)
- chytridiomycota (2)
- cis-regulatory evolution (2)
- click chemistry (2)
- climate adaptation (2)
- climate dynamics (2)
- climate extremes (2)
- climate warming (2)
- co-expression (2)
- co-limitation (2)
- coefficient (2)
- coffee by-products (2)
- cold stress (2)
- collagen (2)
- colon carcinogenesis (2)
- colony viability (2)
- common‐garden experiment (2)
- community model (2)
- community structure (2)
- community theory (2)
- competition–defense trade‐off (2)
- comprehensive analysis (2)
- concepts (2)
- confocal microscopy (2)
- conformational change (2)
- conformational rearrangement (2)
- congeneric species (2)
- conscripts (2)
- constitutive activity (2)
- constraint-based modeling (2)
- converting factor (2)
- copper complex (2)
- copper(II) (2)
- copper-related disorders (2)
- cord blood (2)
- core shell UCNP (2)
- cori cycle (2)
- cortisol (2)
- counting (2)
- coviability analysis (2)
- crop (2)
- crop diversity (2)
- cropping system (2)
- cross-species capture (2)
- cryolithology (2)
- cryptomycota (2)
- cyanobacterial bloom (2)
- cyanobacterial sucrose-phosphatase (2)
- cytochrome c (2)
- cytoplasmic polyadenylation (2)
- cytosine methylation (2)
- cytosolic tRNA thiolation (2)
- cytotoxicity (2)
- dark virus (2)
- data integration (2)
- ddRAD (2)
- de novo genome assembly (2)
- dead Cas9 (2)
- decline (2)
- defense against predation (2)
- defensive symbiosis (2)
- degraded DNA (2)
- dehydration (2)
- demographic noise (2)
- depressive-like behavior (2)
- desiccation (2)
- design of experiment (2)
- developing brain (2)
- developmental canalization (2)
- developmental dyslexia (2)
- developmental plasticity (2)
- diacylglycerol (2)
- dietary patterns (2)
- dietary restriction (2)
- differential expression analysis (2)
- digestive enzymes quantification (2)
- dimerization of 4-nitrothiophenol (2)
- dispersal filtering (2)
- diversity profiles (2)
- dominance effect (2)
- drought stress (2)
- droughts (2)
- drug delivery (2)
- drug metabolism (2)
- drug release (2)
- dynamic equilibrium (2)
- eastern continental Asia (2)
- eavesdropping (2)
- echolocation (2)
- ecophysiology (2)
- ecosystem services provisioning (2)
- education (2)
- effectors (2)
- egg ratio (2)
- eicosapentaenoic acid (2)
- electric fish (2)
- electroencephalography (EEG) (2)
- electron microscopy (2)
- electron paramagnetic resonance (2)
- electronic tool integration (2)
- emotional imagery (2)
- emotions (2)
- endocardium (2)
- endothelial cell (2)
- endotoxin (2)
- energy expenditure (2)
- enrichment experiments (2)
- entropy (2)
- environmental DNA (2)
- environmental genomics (2)
- environmental noise (2)
- enzymology (2)
- epidemiology (2)
- epigenetic variation (2)
- epiphytes (2)
- epithionitrile (2)
- epitope prediction (2)
- epizoochory (2)
- equalizing and stabilizing mechanisms (2)
- error reduction (2)
- establishment (2)
- event coincidence analysis (2)
- evolutionary ecology (2)
- evolutionary rescue (2)
- expansion microscopy (2)
- exploitation (2)
- expression profile (2)
- extinction (2)
- extinction drivers (2)
- extra-cytoplasmic pockets (2)
- extracellular DNA (2)
- extracellular enzymes (2)
- extracellular matrix (2)
- extracellular signaling (2)
- eye-tracking (2)
- facilitation (2)
- fatty acid changes (2)
- feeding (2)
- feeding behaviour (2)
- fence interaction (2)
- fertilization (2)
- fetal origins hypothesis (2)
- fisheries (2)
- fitness gradient (2)
- fitness response (2)
- floating mat (2)
- flooded grasslands (2)
- floral scent (2)
- flow cytometry (2)
- fluorescent probe (2)
- food frequency questionnaire (2)
- food web dynamics (2)
- food webs (2)
- forage availability (2)
- forage gaps (2)
- foraging (2)
- foraging behaviour (2)
- forebrain (2)
- forest management (2)
- forestREplot (2)
- formaldehyde assimilation (2)
- formate dehydrogenase (2)
- fractionation (2)
- fractionation factors (2)
- free zinc (2)
- freshwater algae (2)
- freshwater heterotrophic bacteria (2)
- functional complementation (2)
- functional inhibitors of acid sphin-gomyelinase (2)
- functional response (2)
- functional richness (2)
- fungal diversity (2)
- fungal pathogens (2)
- funktionelle Diversität (2)
- galactolipids (2)
- gamma diversity (2)
- gas-production (2)
- gene delivery (2)
- generalized dissimilarity modelling (2)
- genetic accommodation (2)
- genetic adaptation (2)
- genetic engineering (2)
- genetic rescue (2)
- genetic screen (2)
- genetischer Screen (2)
- genome (2)
- genome scan (2)
- genomic prediction (2)
- geo-bio interaction (2)
- geographic distribution (2)
- glacial maximum (2)
- global and regional change (2)
- glucocorticoid receptor (2)
- glucose (2)
- glucosinolate hydrolysis (2)
- glutathione (2)
- glycine cleavage system (2)
- governance (2)
- grazing (2)
- gross primary production (2)
- groundwater (2)
- groundwater recharge (2)
- growth behavior (2)
- growth restriction (2)
- guard cell (2)
- gut microbiota (2)
- gwas (2)
- habitat (2)
- habitat selection (2)
- habitat use (2)
- habituation (2)
- heart regeneration (2)
- heat (2)
- heat shock protein (2)
- heliozoa (2)
- hepcidin-25 (2)
- heterocyclic aromatic amine (2)
- hilly loes plateau (2)
- histone modification (2)
- holding capability (2)
- holding isometric muscle action (HIMA) (2)
- holocene (2)
- honey bee (2)
- honey bees (2)
- hoverflies (2)
- human aldehyde oxidase (2)
- human endotoxemia (2)
- human excised skin (2)
- human introduction (2)
- human-wildlife conflict (2)
- hydrogels (2)
- hyena (2)
- hyperoxia (2)
- hypertension (2)
- hyperthermia (2)
- immunoassay (2)
- immunogenicity (2)
- importin (2)
- in silico (2)
- in vitro intestinal model (2)
- in-vitro-synthesis (2)
- inbreeding depression (2)
- indirect facilitation (2)
- individual based modeling (2)
- individual-based modeling (2)
- infancy (2)
- infiltration (2)
- inhibition (2)
- inner-mongolia (2)
- integrated assessments (2)
- inter-brain synchronization (2)
- inter-muscle-brain synchronization (2)
- interpersonal muscle action (2)
- interspecific interactions (2)
- intestinal mucins (2)
- intestinal zinc resorption (2)
- intra-organ-communication (2)
- intraguild predation (2)
- intraspecific variation (2)
- invasibility (2)
- invasion success (2)
- invasive species (2)
- ion channel (2)
- ion mobility spectrometry (2)
- ion transport (2)
- ion-exchange chromatography (2)
- ionic liquid (2)
- ionic strength (2)
- ionogel (2)
- iron-sulfur clusters (2)
- island biogeography (2)
- island disharmony (2)
- island syndromes (2)
- islands (2)
- isomerization (2)
- jasmonate (2)
- kelp (2)
- l-cysteine desulfurase (2)
- lake monitoring (2)
- lake periphyton (2)
- land sharing vs. land sparing (2)
- land use change (2)
- land-use change (2)
- landscape diversity (2)
- landscape generator (2)
- landscape homogenization (2)
- late pleistocene (2)
- later health (2)
- lichens (2)
- life cycle (2)
- life‐history traits (2)
- ligand exchange (2)
- light adaptation (2)
- light variability (2)
- limits (2)
- lipid classes (2)
- lipid limitation thresholds (2)
- lipid membranes (2)
- lipid–lipid interactions (2)
- lipoplexes (2)
- loci (2)
- lokale Anpassung (2)
- longitudinal (2)
- low-energy electrons (2)
- luminescence (2)
- lysosomal hydrolases (2)
- lysosomal storage disorders (2)
- lysosome (2)
- mRNA degradation (2)
- magnitude estimation (2)
- maintenance (2)
- maintenance of functional diversity (2)
- maintenance of genomic integrity (2)
- major histocompatibility complex (2)
- male Daphnia (2)
- maltooligosaccharides (2)
- mammalian-cells (2)
- manganese (2)
- manual muscle test (2)
- many-to-one genotype–phenotype map (2)
- mapping (2)
- mass conservation (2)
- mass index (2)
- mate-pairs (2)
- mathematical modelling (2)
- mathematical precursor (2)
- maturation (2)
- meal timing (2)
- mechanical (2)
- mechanical strength (2)
- mechanisms (2)
- mechanomyography (MMG) (2)
- mediated delivery (2)
- membrane biophysics (2)
- membrane fusion (2)
- membrane microdomains (2)
- membrane protein (2)
- membrane stabilization (2)
- memory (2)
- mercaptocarboxylic acids (2)
- mesenchymal stem cells (2)
- mesophyll cell (2)
- messenger-rna polyadenylation (2)
- meta-analysis (2)
- metabolic syndrome (2)
- metabolic-profiling (2)
- metabolische Netzwerke (2)
- metabolite (2)
- metagenome (2)
- metagenomic analysis (2)
- metagenomics 2.0 (2)
- metal complex (2)
- metal peptide (2)
- metallopeptide (2)
- metalloprotein (2)
- methane oxidation (2)
- methylotrophy (2)
- miRNAs (2)
- microarrays (2)
- microbial activity (2)
- microbial communities (2)
- microbiota (2)
- microclimate (2)
- microcomputed tomography (2)
- microeukaryotes (2)
- microfluidics (2)
- microstructure (2)
- microtubule-organization (2)
- mitochondrial genome (mtDNA) (2)
- mitogenomes (2)
- mixed cultures (2)
- model (2)
- model integration (2)
- model limitations (2)
- modeling (2)
- models (2)
- modern coexistence theory (2)
- mojave desert (2)
- molecular biology (2)
- molecular dynamics simulations (2)
- molecular evolution (2)
- molecular species identification (2)
- molybdoenzyme (2)
- molybdopterin synthase (2)
- morphology (2)
- movement speed (2)
- mucus layer (2)
- multidrug resistance (2)
- multiple sclerosis (2)
- multishell (2)
- multivalence (2)
- multi‐ year flooding cycle (2)
- muscle (2)
- musicality (2)
- mutation (2)
- mutual information (2)
- mycotoxins (2)
- myocardial infarction (2)
- myocardium (2)
- myoglobin (2)
- n-alkanes (2)
- n-oxide reductase (2)
- nachhaltige Landnutzung (2)
- nanoelectrodes (2)
- nanogels (2)
- nanostructure (2)
- native American ancestry (2)
- necrobiome (2)
- neophilia (2)
- neophobia (2)
- net primary productivity (2)
- network analysis (2)
- network reconstruction (2)
- neurodegeneration (2)
- neurodegenerative diseases (2)
- neuroendocrine (2)
- neuromuscular adaptation (2)
- niche and fitness differences (2)
- niche width (2)
- nickel (2)
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- nitrogen (2)
- nitrogen fixation (2)
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Institute
- Institut für Biochemie und Biologie (1806)
- Institut für Ernährungswissenschaft (242)
- Mathematisch-Naturwissenschaftliche Fakultät (120)
- Extern (81)
- Institut für Chemie (55)
- Institut für Geowissenschaften (38)
- Institut für Physik und Astronomie (35)
- Institut für Umweltwissenschaften und Geographie (31)
- Department Sport- und Gesundheitswissenschaften (14)
- Fakultät für Gesundheitswissenschaften (10)
The multidrug and toxic compounds extrusion (MATE) family includes hundreds of functionally uncharacterised proteins from bacteria and all eukaryotic kingdoms except the animal kingdom, that function as drug/toxin::Na<sup>+ or H<sup>+ antiporters. In Arabidopsis thaliana the MATE family comprises 56 members, one of which is NIC2 (Novel Ion Carrier 2). Using heterologous expression systems including Escherichia coli and Saccharomyces cerevisiae, and the homologous expression system of Arabidopsis thaliana, the functional characterisation of NIC2 was performed. It has been demonstrated that NIC2 confers resistance of E. coli towards the chemically diverse compounds such as tetraethylammonium chloride (TEACl), tetramethylammonium chloride (TMACl) and a toxic analogue of indole-3-acetic acid, 5-fluoro-indole-acetic acid (F-IAA). Therefore, NIC2 may be able to transport a broad range of drug and toxic compounds. In wild-type yeast the expression of NIC2 increased the tolerance towards lithium and sodium, but not towards potassium and calcium. In A. thaliana, the overexpression of NIC2 led to strong phenotypic changes. Under normal growth condtions overexpression caused an extremely bushy phenotype with no apical dominance but an enhanced number of lateral flowering shoots. The amount of rossette leaves and flowers with accompanying siliques were also much higher than in wild-type plants and the senescence occurred earlier in the transgenic plants. In contrast, RNA interference (RNAi) used to silence NIC2 expression, induced early flower stalk development and flowering compared with wild-type plants. In additon, the main flower stalks were not able to grow vertically, but instead had a strong tendency to bend towards the ground. While NIC2 RNAi seedlings produced many lateral roots outgrowing from the primary root and the root-shoot junction, NIC2 overexpression seedlings displayed longer primary roots that were characterised by a 2 to 4 h delay in the gravitropic response. In addition, these lines exhibited an enhanced resistance to exogenously applied auxins, i.e. indole-3-acetic acid (IAA) and indole-3-butyric acid (IBA) when compared with the wild-type roots. Based on these results, it is suggested that the NIC2 overexpression and NIC2 RNAi phenotypes were due to decreased or increased levels of auxin, respectively. The ProNIC2:GUS fusion gene revealed that NIC2 is expressed in the stele of the elongation zone, in the lateral root cap, in new lateral root primordia, and in pericycle cells of the root system. In the vascular tissue of rosette leaves and inflorescence stems, the expression was observed in the xylem parenchyma cells, while in siliques it was also in vascular tissue, but as well in the dehiscence and abscission zones. The organ- and tissue-specific expression sites of NIC2 correlate with the sites of auxin action in mature Arabidopsis plants. Further experiments using ProNIC2:GUS indicated that NIC2 is an auxin-inducible gene. Additionally, during the gravitropic response when an endogenous auxin gradient across the root tip forms, the GUS activity pattern of the ProNIC2:GUS fusion gene markedly changed at the upper side of the root tip, while at the lower side stayed unchanged. Finally, at the subcellular level NIC2-GFP fusion protein localised in the peroxisomes of Nicotana tabacum BY2 protoplasts. Considering the experimental results, it is proposed that the hypothetical function of NIC2 is the efflux transport which takes part in the auxin homeostasis in plant tissues probably by removing auxin conjugates from the cytoplasm into peroxisomes.
The protection of species is one major focus in conservation biology. The basis for any management concept is the knowledge of the species autecology. In my thesis, I studied the life-history traits and population dynamics of the endangered Lesser Spotted Woodpecker (Picoides minor) in Central Europe. Here, I combine a range of approaches, from empirical investigations of a Lesser Spotted Woodpecker population in the Taunus low mountain range in Germany, the analysis of empirical data and the development of an individual-based stochastic model simulating the population dynamics. In the field studies I collected basic demographic data of reproductive success and mortality. Moreover, breeding biology and behaviour were investigated in detail. My results showed a significant decrease of the reproductive success with later timing of breeding, caused by deterioration in food supply. Moreover, mate fidelity was of benefit, since pairs composed of individuals that bred together the previous year started earlier with egg laying and obtained a higher reproductive success. Both sexes were involved in parental care, but the care was only shared equally during incubation and the early nestling stage. In the late nestling stage, parental care strategies differed between sexes: Females considerably decreased feeding rate with number of nestlings and even completely deserted small broods. Males fed their nestlings irrespective of brood size and compensated for the females absence. The organisation of parental care in the Lesser Spotted Woodpecker is discussed to provide the possibility for females to mate with two males with separate nests and indeed, polyandry was confirmed. To investigate the influence of the observed flexibility in the social mating system on the population persistence, a stochastic individual-based model simulating the population dynamics of the Lesser Spotted Woodpecker was developed, based on empirical results. However, pre-breeding survival rates could not be obtained empirically and I present in this thesis a pattern-oriented modelling approach to estimate pre-breeding survival rates by comparing simulation results with empirical pattern of population structure and reproductive success on population level. Here, I estimated the pre-breeding survival for two Lesser Spotted Woodpecker populations on different latitudes to test the reliability of the results. Finally, I used the same simulation model to investigate the effect of flexibility in the mating system on the persistence of the population. With increasing rate of polyandry in the population, the persistence increased and even low rates of polyandry had a strong influence. Even when presuming only a low polyandry rate and costs of polyandry in terms of higher mortality and lower reproductive success for the secondary male, the positive effect of polyandry on the persistence of the population was still strong. This thesis greatly helped to increase the knowledge of the autecology of an endangered woodpecker species. Beyond the relevance for the species, I could demonstrate here that in general flexibility in mating systems are buffer mechanisms and reduce the impact of environmental and demographic noise.
During this PhD project three technical platforms were either improved or newly established in order to identify interesting genes involved in SNF, validate their expression and functionally characterise them. An existing 5.6K cDNA array (Colebatch et al., 2004) was extended to produce the 9.6K LjNEST array, while a second array, the 11.6K LjKDRI array, was also produced. Furthermore, the protocol for array hybridisation was substantially improved (Ott et al., in press). After functional classification of all clones according to the MIPS database and annotation of their corresponding tentative consensus sequence (TIGR) these cDNA arrays were used by several international collaborators and by our group (Krusell et al., 2005; in press). To confirm results obtained from the cDNA array analysis different sets of cDNA pools were generated that facilitate rapid qRT-PCR analysis of candidate gene expression. As stable transformation of Lotus japonicus takes several months, an Agrobacterium rhizogenes transformation system was established in the lab and growth conditions for screening transformants for symbiotic phenotypes were improved. These platforms enable us to identify genes, validate their expression and functionally characterise them in the minimum of time. The resources that I helped to establish, were used in collaboration with other people to characterise several genes like the potassium transporter LjKup and the sulphate transporter LjSst1, that were transcriptionally induced in nodules compared to uninfected roots, in more detail (Desbrosses et al., 2004; Krusell et al., 2005). Another gene that was studied in detail was LjAox1. This gene was identified during cDNA array experiments and detailed expression analysis revealed a strong and early induction of the gene during nodulation with high expression in young nodules which declines with the age of the nodule. Therefore, LjAox1 is an early nodulin. Promoter:gus fusions revealed an LjAox1 expression around the nodule endodermis. The physiological role of LjAox1 is currently being persued via RNAi. Using RNA interference, the synthesis of all symbiotic leghemoglobins was silenced simultaneously in Lotus japonicus. As a result, growth of LbRNAi lines was severely inhibited compared to wild-type plants when plants were grown under symbiotic conditions in the absence of mineral nitrogen. The nodules of these plants were arrested in growth 14 post inoculation and lacked the characteristic pinkish colour. Growing these transgenic plants in conditions where reduced nitrogen is available for the plant led to normal plant growth and development. This demonstrates that leghemoglobins are not required for plant development per se, and proves for the first time that leghemoglobins are indispensable for symbiotic nitrogen fixation. Absence of leghemoglobins in LbRNAi nodules led to significant increases in free-oxygen concentrations throughout the nodules, a decrease in energy status as reflected by the ATP/ADP ratio, and an absence of the bacterial nitrogenase protein. The bacterial population within nodules of LbRNAi plants was slightly reduced. Alterations of plant nitrogen and carbon metabolism in LbRNAi nodules was reflected in changes in amino acid composition and starch deposition (Ott et al., 2005). These data provide strong evidence that nodule leghemoglobins function as oxygen transporters that facilitate high flux rates of oxygen to the sites of respiration at low free oxygen concentrations within the infected cells.
Vitamin E : elucidation of the mechanism of side chain degradation and gene regulatory functions
(2005)
For more than 80 years vitamin E has been in the focus of scientific research. Most of the progress concerning non-antioxidant functions, nevertheless, has only arisen from publications during the last decade. Most recently, the metabolic pathway of vitamin E has been almost completely elucidated. Vitamin E is metabolized by truncation of its side chain. The initial step of an omega-hydroxylation is carried out by cytochromes P450 (CYPs). This was evidenced by the inhibition of the metabolism of alpha-tocopherol by ketoconozole, an inhibitor of CYP3A expression, whereas rifampicin, an inducer of CYP3A expression increased the metabolism of alpha-tocopherol. Although the degradation pathway is identical for all tocopherols and tocotrienols, there is a marked difference in the amount of the release of metabolites from the individual vitamin E forms in cell culture as well as in experimental animals and in humans. Recent findings not only proposed an CYP3A4-mediated degradation of vitamin E but also suggested an induction of the metabolizing enzymes by vitamin E itself. In order to investigate how vitamin E is able to influence the expression of metabolizing enzymes like CYP3A4, a pregnane X receptor (PXR)-based reporter gene assay was chosen. PXR is a nuclear receptor which regulates the transcription of genes, e.g., CYP3A4, by binding to specific DNA response elements. And indeed, as shown here, vitamin E is able to influence the expression of CYP3A via PXR in an in vitro reporter gene assay. Tocotrienols showed the highest activity followed by delta- and alpha-tocopherol. An up-regulation of Cyp3a11 mRNA, the murine homolog of the human CYP3A4, could also be confirmed in an animal experiment. The PXR-mediated change in gene expression displayed the first evidence of a direct transcriptional activity of vitamin E. PXR regulates the expression of genes involved in xenobiotic detoxification, including oxidation, conjugation, and transport. CYP3A, e.g., is involved in the oxidative metabolism of numerous currently used drugs. This opens a discussion of possible side effects of vitamin E, but the extent to which supranutritional doses of vitamin E modulate these pathways in humans has yet to be determined. Additionally, as there is arising evidence that vitamin E's essentiality is more likely to be based on gene regulation than on antioxidant functions, it appeared necessary to further investigate the ability of vitamin E to influence gene expression. Mice were divided in three groups with diets (i) deficient in alpha-tocopherol, (ii) adequate in alpha-tocopherol supply and (iii) with a supranutritional dosage of alpha-tocopherol. After three months, half of each group was supplemented via a gastric tube with a supranutritional dosage of gamma-tocotrienol per day for 7 days. Livers were analyzed for vitamin E content and liver RNA was prepared for hybridization using cDNA array and oligonucleotide array technology. A significant change in gene expression was observed by alpha-tocopherol but not by gamma-tocotrienol and only using the oligonucleotide array but not using the cDNA array. The latter effect is most probably due to the limited number of genes represented on a cDNA array, the lacking gamma-tocotrienol effect is obviously caused by a rapid degradation, which might prevent bioefficacy of gamma-tocotrienol. Alpha-tocopherol changed the expression of various genes. The most striking observation was an up-regulation of genes, which code for proteins involved in synaptic transmitter release and calcium signal transduction. Synapsin, synaptotagmin, synaptophysin, synaptobrevin, RAB3A, complexin 1, Snap25, ionotropic glutamate receptors (alpha 2 and zeta 1) were shown to be up-regulated in the supranutritional group compared to the deficient group. The up-regulation of synaptic genes shown in this work are not only supported by the strong concentration of genes which all are involved in the process of vesicular transport of neurotransmitters, but were also confirmed by a recent publication. However, a confirmation by real time PCR in neuronal tissue like brain is now required to explain the effect of vitamin E on neurological functionality. The change in expression of genes coding for synaptic proteins by vitamin E is of principal interest thus far, since the only human disease directly originating from an inadequate vitamin E status is ataxia with isolated vitamin E deficiency. Therefore, with the results of this work, an explanation for the observed neurological symptoms associated with vitamin E deficiency can be presented for the first time.
Für ein tiefergehendes Verständnis von Entwicklung und Funktion der quergestreiften Muskulatur ist eine Betrachtung der am Aufbau der Myofibrillen, den kontraktilen Organellen, beteiligten Proteine essentiell. Die vorliegende Arbeit beschäftigt sich mit Myomesin, einem Protein der sarkomeren M-Bande. Zunächst wurde die cDNA des humanen Myomesins vollständig kloniert, sequenziert und nachfolgend die komplette Größe der aminoterminalen Kopfdomäne bestimmt. Es konnte gezeigt werden, daß Myomesin in vitro mit den Domänen 1 und 12 an Myosin bindet. Die muskelspezifische Isoform der Kreatinkinase bindet an die Domänen 7 und 8. Stimulations- und Inhibitionsexperimente belegen, daß Myomesin an Serin 618 in vivo durch die Proteinkinase A phosphoryliert wird und daß diese Phosphorylierung durch Aktivierung beta2-adrenerger Rezeptoren stimulierbar ist. In Muskelgewebeproben von Patienten, die an der Hypertrophen Kardiomyopathie, einer genetisch bedingten Herzmuskelkrankheit, erkrankt sind, konnte mit einem neu hergestellten phosphorylierungsabhängigen Antikörper eine Verminderung der Menge phosphorylierten Myomesins nachgewiesen werden. Mögliche Ursachen werden diskutiert. Myomesin bildet Dimere, wie durch hefegenetische und biochemische Experimente gezeigt werden konnte. Die Dimerisierung von Myomesin könnte eine zentrale Rolle für den Einbau der Myosinfilamente in die naszierende Myofibrille haben. Anhand der gewonnenen Daten wurde ein verbessertes Modell der zentralen M-Bande erstellt.
Even though the structure of the plant cell wall is by and large quite well characterized, its synthesis and regulation remains largely obscure. However, it is accepted that the building blocks of the polysaccharidic part of the plant cell wall are nucleotide sugars. Thus to gain more insight into the cell wall biosynthesis, in the first part of this thesis, plant genes possibly involved in the nucleotide sugar interconversion pathway were identified using a bioinformatics approach and characterized in plants, mainly in Arabidopsis. For the computational identification profile hidden markov models were extracted from the Pfam and TIGR databases. Mainly with these, plant genes were identified facilitating the “hmmer” program. Several gene families were identified and three were further characterized, the UDP-rhamnose synthase (RHM), UDP-glucuronic acid epimerase (GAE) and the myo-inositol oxygenase (MIOX) families. For the three-membered RHM family relative ubiquitous expression was shown using variuos methods. For one of these genes, RHM2, T-DNA lines could be obtained. Moreover, the transcription of the whole family was downregulated facilitating an RNAi approach. In both cases a alteration of cell wall typic polysaccharides and developmental changes could be shown. In the case of the rhm2 mutant these were restricted to the seed or the seed mucilage, whereas the RNAi plants showed profound changes in the whole plant. In the case of the six-membered GAE family, the gene expressed to the highest level (GAE6) was cloned, expressed heterologously and its function was characterized. Thus, it could be shown that GAE6 encodes for an enzyme responsible for the conversion of UDP-glucuronic acid to UDP-galacturonic acid. However, a change in transcript level of variuos GAE family members achieved by T-DNA insertions (gae2, gae5, gae6), overexpression (GAE6) or an RNAi approach, targeting the whole family, did not reveal any robust changes in the cell wall. Contrary to the other two families the MIOX gene family had to be identified using a BLAST based approach due to the lack of enough suitable candidate genes for building a hidden markov model. An initial bioinformatic characterization was performed which will lead to further insights into this pathway. In total it was possible to identify the two gene families which are involved in the synthesis of the two pectin backbone sugars galacturonic acid and rhamnose. Moreover with the identification of the MIOX genes a genefamily, important for the supply of nucleotide sugar precursors was identified. In a second part of this thesis publicly available microarray datasets were analyzed with respect to co-responsive behavior of transcripts on a global basis using nearly 10,000 genes. The data has been made available to the community in form of a database providing additional statistical and visualization tools (http://csbdb.mpimp-golm.mpg.de). Using the framework of the database to identify nucleotide sugar converting genes indicated that co-response might be used for identification of novel genes involved in cell wall synthesis based on already known genes.
Durch die anthropogene Nutzung sind viele Auen in Mitteleuropa verändert worden, wobei insbesondere die Retentionsflächen stark verringert wurden. Während Auen seit längerem im Fokus der wissenschaftlichen Bearbeitung stehen, gibt es bisher große Wissensdefizite in der Frage der Auenreaktivierungen. Zum einen sind derartige Projekte bisher kaum verwirklicht und zum anderen ist ein langfristiges Monitoring notwendig, um die Anpassung von Biozönosen an die veränderten Standortbedingungen beobachten zu können. Um die Folgen derartiger Eingriffe zu analysieren, bieten sich computergestützte Modellierungen der Landschaftsentwicklung an, wie sie in der vorliegenden Arbeit verwirklicht wurden. Ziel der Arbeit war, mit Hilfe eines Geografischen Informationssystems (GIS) das Entwicklungspotenzial der Landschaft bei verschiedenen Rückdeichungsvarianten auf der Ebene der Biotoptypen darzustellen. Dabei ging es nicht um die Erstellung eines allgemein gültigen Auenmodells sondern um die Erarbeitung eines Modells für einen konkreten Anwendungsfall. Der erarbeitete Ansatz sollte zudem für die landschaftsplanerische Praxis geeignet sein. Als Beispielgebiete wurden Flächen an der Mittleren Elbe bei Rogätz und Sandau, beide im nördlichen Teil von Sachsen-Anhalt, ausgewählt. Die vorliegende Arbeit gliedert sich in zwei Teile. Im ersten Teil werden Erhebungen und Auswertungen als Grundlage der Modellentwicklung dargestellt. Dazu wurden die Biotoptypen der Beispielgebiete flächendeckend erhoben und mit punktuellen Vegetationserhebungen ergänzt. Aus dem Forschungsprojekt "Rückgewinnung von Retentionsflächen und Altauenreaktivierung an der Mittleren Elbe in Sachsen-Anhalt" des Bundesministeriums für Bildung und Forschung (BMBF) standen standortökologische Daten der Hydrologie und Bodenkunde zur Verfügung. Ziel der Auswertung war, Schlüsselfaktoren für Hydrologie und Bodenbedingungen innerhalb der rezenten Aue zu identifizieren, die zur Ausprägung bestimmter Biotoptypen führen. Im zweiten Teil der Arbeit wurde ein Modell für Biotoptypenpotenziale auf den geplanten Rück–deichungsflächen entwickelt. Das Modell bearbeitet die Datenbank der verwendeten GIS-Dateien, die auf Daten zum Bestand beruht und um solche der Prognose der Standortökologie (Hydrologie und Boden) im Rückdeichungsfalle aus dem BMBF-Projekt erweitert wurde. Weitere Voraussetzung für die Modellierung war die Erarbeitung von Leitbildern, in denen unterschiedliche Nutzungsszenarios für die Landschaft nach Deichrückverlegung hypothetisch festgelegt wurden. Insbesondere die Nutzungsintensität wurde variiert, von einer Variante intensiver land- und forstwirtschaftlicher Nutzung über sogenannte integrierte Entwicklungsziele aus dem BMBF-Projekt bis hin zu einer Variante der Naturschutznutzung. Zusätzlich wurde eine zukünftige Potentielle Natürliche Vegetation modelliert. Eine Überprüfung des Modell fand für den Raum der rezenten Aue in der intensiven Nutzungsvariante statt, die der gegenwärtigen Nutzung am nächsten kommt. Werden Informationen des Bestandsbiotoptyps als Korrekturgröße in das Modell einbezogen, konnte für viele Biotoptypen eine Trefferquote von über 90 % erreicht werden. Bei flächenmäßig weniger bedeutenden Bio–toptypen lag dieser Wert aufgrund der schmaleren Datenbasis zwischen 20 und 40 %. Als Ergebnis liegt für unterschiedliche Deichvarianten und Leitbilder in den Beispielgebieten die Landschaftsentwicklung als Biotoppotenzial vor. Als eine vereinfachte Regionalisierung der punktuellen Vegetationsdaten wurde im Modell geprüft, inwieweit die modellierten Biotopflächen der Charakteristik der pflanzensoziologischen Aufnahmen aus der rezenten Aue entsprechen. In dem Falle wurde die Pflanzengesellschaft der jeweiligen ökologisch im Rahmen der Untersuchung einheitlichen Flächeneinheit zugeordnet. Anteilig lässt sich damit die Biotopprognosefläche pflanzensoziologisch konkretisieren. Die vorliegende Arbeit gehört zu den bisher wenigen Arbeiten, die sich mit den Folgen von Auenreaktivierung auf die Entwicklung der Landschaft auseinandersetzen. Sie zeigt eine Möglichkeit auf, Prognosemodelle für Biotoptypen und Vegetation anhand begrenzter Felduntersuchungen zu entwerfen. Derartige Modelle können zum Verständnis von Eingriffen in den Naturhaushalt, wie sie die Deichrückverlegungen darstellen, beitragen und eine Folgenabschätzung unterstützen.
Im Mittelpunkt dieser Arbeit standen Signaltransduktionsprozesse in den Strukturen der Kraftübertragung quergestreifter Muskelzellen, d. h. in den Costameren (Zell-Matrix-Kontakten) und den Glanzstreifen (Zell-Zell-Kontakten der Kardiomyozyten).Es ließ sich zeigen, dass sich die Morphologie der Zell-Matrix-Kontakte während der Differenzierung von Skelettmuskelzellen dramatisch ändert, was mit einer veränderten Proteinzusammensetzung einhergeht. Immunfluoreszenz-Analysen von Skelettmuskelzellen verschiedener Differenzierungsstadien implizieren, dass die Signalwege, welche die Dynamik der Fokalkontakte in Nichtmuskelzellen bestimmen, nur für frühe Stadien der Muskeldifferenzierung Relevanz haben können. Ausgehend von diesem Befund wurde begonnen, noch unbekannte Signalwege zu identifizieren, welche die Ausbildung von Costameren kontrollieren: In den Vorläuferstrukturen der Costamere gelang es, eine transiente Interaktion der Proteine Paxillin und Ponsin zu identifizieren. Biochemische Untersuchungen legen nahe, dass Ponsin über eine Skelettmuskel-spezifische Insertion im Carboxyterminus das Adapterprotein Nck2 in diesen Komplex rekrutiert. Es wird vorgeschlagen, dass die drei Proteine einen ternären Signalkomplex bilden, der die Umbauvorgänge der Zell-Matrix-Kontakte kontrolliert und dessen Aktivität von mitogen activated protein kinases (MAPK) reguliert wird.Die Anpassungsvorgänge der Strukturen der Kraftübertragung an pathologische Situtation (Kardiomyopathien) in der adulten quergestreiften Muskulatur wurden ausgehend von einem zweiten Protein, dem muscle LIM protein (MLP), untersucht. Es konnte gezeigt werden, dass ein mutiertes MLP-Protein, das im Menschen eine hypertrophe Kardiomyopathie (HCM) auslöst, strukturelle Defekte aufweist und weniger stabil ist. Weiterhin zeigte dieses mutierte Protein eine verringerte Bindungsfähigkeit an die beiden Liganden N-RAP und alpha-Actinin. Die molekulare Grundlage der HCM-verursachenden Mutationen im MLP-Gen könnte folglich eine Veränderung der Homöostase im ternären Komplex MLP – N-RAP – alpha-Actinin sein. Die Expressionsdaten eines neu generierten monoklonalen MLP-Antikörpers deuten darauf hin, dass die Funktionen des MLP nicht nur für die Integrität des Myokards, sondern auch für die der Skelettmuskulatur notwendig sind.
Für das Verständnis der Strukturbildung bei Proteinen ist es wichtig, allgemein geltende Prinzipien der Stabilität und Faltung zu verstehen. Bisher wurde viel Arbeit in die Erörterung von Gesetzmäßigkeiten zu den Faltungseigenschaften von globulären Proteinen investiert. Die große Proteinklasse der solenoiden Proteine, zu denen z. B. Leucine-Rich Repeat- (LRR-) oder Ankyrin-Proteine gehören, wurde dahingegen noch wenig untersucht. Die Proteine dieser Klasse sind durch einen stapelförmigen Aufbau von sich wiederholenden typischen Sequenzeinheiten gekennzeichnet, was in der Ausbildung einer elongierten Tertiärstruktur resultiert. In der vorliegenden Arbeit sollte versucht werden, die Stabilität und Faltung eines LRR-Proteins mittels verschiedener biophysikalischer Methoden zu charakterisieren. Als Untersuchungsobjekt diente die für die Infektion ausreichende zentrale LRR-Domäne des Invasionsproteins Internalin B (InlB241) des Bakteriums Listeria monocytogenes. Des weiteren sollten die Integrität und die Stabilitäts- und Faltungseigenschaften der sogenannten Internalin-Domäne (InlB321) untersucht werden. Hierbei handelt es sich um die bei allen Mitgliedern der Internalinfamilie vorkommende Domäne, welche aus einer direkten Fusion des C-terminalen Endes der LRR-Domäne mit einer Immunglobulin (Ig)-ähnlichen Domäne besteht. Von beiden Konstrukten konnte eine vollständige thermodynamische Charakterisierung, mit Hilfe von chemisch- bzw. thermisch-induzierten Faltungs- und Entfaltungsübergängen durchgeführt werden. Sowohl InlB241 als auch InlB321 zeigen einen reversiblen und kooperativen Verlauf der chemisch-induzierten Gleichgewichtsübergänge, was die Anwendung eines Zweizustandsmodells zur Beschreibung der Daten erlaubte. Die zusätzliche Ig-ähnliche Domäne im InlB321 resultierte im Vergleich zum InlB241 in einer Erhöhung der freien Enthalpie der Entfaltung (8.8 kcal/mol im Vergleich zu 4.7 kcal/mol). Diese Stabilitätszunahme äußerte sich sowohl in einer Verschiebung des Übergangsmittelpunktes zu höheren Guanidiniumchlorid-Konzentrationen als auch in einer Erhöhung der Kooperativität des Gleichgewichtsübergangs (9.7 kcal/mol/M im Vergleich zu 7.1 kcal/mol/M). Diese Beobachtungen zeigen dass die einzelnen Sequenzeinheiten der LRR-Domäne nicht unabhängig voneinander falten und dass die Ig-ähnliche Domäne, obwohl sie nicht direkt mit dem Wirtszellrezeptor während der Invasion interagiert, eine kritische Rolle für die <i style='mso-bidi-font-style:normal'>in vivo Stabilität des Internalin B spielt. Des weiteren spiegelt die Kooperativität des Übergangs die Integrität der Internalin-Domäne wieder und deutet darauf hin, dass bei beiden Proteinen keine Intermediate vorliegen. Kinetische Messungen über Tryptophanfluoreszenz und Fern-UV<span style='color:red'> </span>Circulardichroismus deuteten auf die Existenz eines relativ stabilen Intermediates auf dem Faltungsweg der LRR-Domäne hin. Faltungskinetiken aus einem in pH 2 denaturierten Zustand zeigten ein reversibles Verhalten und verliefen über ein Intermediat. Eine Erhöhung der Salzkonzentration des sauer-denaturierten Proteins führte zu einer Kompaktierung der entfalteten Struktur und resultierte im Übergang zu einem alternativ gefalteten Zustand. Bei der Internalin-Domäne deuteten kinetische Messungen des Fluoreszenz- und Fern-UV Circulardichroismus-Signals während der Entfaltung möglicherweise auf die Präsenz von zwei Prozessen hin. Der erste langsame Entfaltungsprozess kurz nach dem Übergangsmittelpunkt zeigte eine starke Abhängigkeit von der Temperatur, während der zweite schnellere Prozess der Entfaltung stärker von der Guanidiniumchlorid-Konzentration abhing. Renaturierungskinetiken zeigten das Auftreten von mindestens einem Faltungsintermediat. Kinetische Daten aus Doppelsprungexperimenten lieferten für die Erklärung der langsamen Faltungsphase zunächst keinen Hinweis auf dass Vorliegen einer Prolinisomerisierungsreaktion. Die vollständige Amplitude während der Renaturierung konnte nicht detektiert werden, weswegen von einer zweiten schnellen Phase im Submillisekundenbereich ausgegangen werden kann. Die Ergebnisse der Faltungskinetiken zeigen, dass die InlB-Konstrukte als Modelle für die Untersuchung der Faltung von Solenoidproteinen verwendet werden können.<span lang=EN-GB style='mso-ansi-language: EN-GB'><o:p></o:p></span>
Charakterisierung von ausgewählten Protein-Phosphatasen und MATE-Proteinen aus Arabidopsis thaliana
(2004)
Im ersten Teil der vorliegenden Arbeit wurde die Genexpression der Protein Phosphatase-gene TOPP1, TOPP2, TOPP5, STH1 und STH2 analysiert. Alle fünf ausgewählten Gene kodieren für PP des PP1/PP2A-Typs. Es wurde untersucht, ob homologen PP-Isoformen individuelle Expressionsmuster zugewiesen werden konnten. Besonderes Augenmerk richtete sich dabei auf die Expression von PP-Genen in den Schließzellen von A. thaliana. In mehreren Inhibitorstudien wurde beschrieben, dass PP1/PP2A-Proteine eine wichtige Rolle in der Signaltransduktion pflanzlicher Schließzellen spielen. Bisher konnte allerdings noch keine der entsprechenden katalytischen Untereinheiten auf molekularer Ebene identifiziert werden. Im Rahmen dieser Arbeit wurde zum ersten Mal nachgewiesen, dass mit TOPP1 ein Protein des PP1-Typs präferenziell in den Schließzellen von A. thaliana exprimiert wird. Ein Vergleich der Genexpression von TOPP1, TOPP2 und TOPP5 zeigte für die drei homologen Gene sehr Isoform-spezifische Expressionsmuster. Dies war ein deutlicher Hinweis, dass diese eng verwandten PP trotz großer Übereinstimmung auf Aminosäureebene vermutlich unterschiedliche Funktionen in planta haben. Die Untersuchung der Genexpression von STH1 und STH2 zeigte, dass die fast identischen Proteine zum Teil in unterschiedlichen Geweben vorkommen. Die Transkripte der beiden Gene, welche eine eigene Untergruppe von PP2A-verwandten Sequenzen bilden, konnten aus EF isoliert werden. Der in dieser Arbeit entwickelte Screeningansatz ermöglichte es, die sehr ähnlichen cDNA-Fragmente eindeutig voneinander zu unterscheiden. Die gefundenen Isoform-spezifischen Expressionsmuster waren ein deutlicher Hinweis auf unterschiedliche Funktionen in planta. Zur weiteren Untersuchung der PP-Funktionen in planta wurden Pflanzen mit veränderter Genaktivität von TOPP2 oder STH1 untersucht. In Pflanzen mit RNAi-vermittelter Reduktion des TOPP2-Transkriptgehalts ließ sich ein deutlich verändertes Blattwachstum beobachten. Die eingerollten oder asymmetrisch entwickelten Blätter waren vermutlich ein Hinweis, dass diese PP1-Isoform auch in A. thaliana eine Rolle bei der Zellteilung spielt. Für TOPP2-Expression in Hefen wurde diese Funktion schon nachgewiesen. Die Analyse von Insertions-mutanten mit T-DNA Insertionen in beiden STH1-Allelen waren neben den Expressions-studien ein weiterer Hinweis, dass sich STH1 nicht funktionell durch STH2 ersetzen lässt. Die Experimente in dieser Arbeit zeigten, dass das Fehlen der STH1-Genaktivität zu einem deutlichen Blattphänotyp mit gezahnten Blatträndern führte. Für STH2-Insertionsmutanten wurde dieses veränderte Wachstum nicht beschrieben. Im zweiten Teil dieser Arbeit wurde das Gen NIC1, welches für ein MATE-Membranprotein kodiert, identifiziert und charakterisiert. Die Sequenzierung des Genoms von A. thaliana hatte gezeigt, dass mindestens 56 MATE-Gene in dieser Pflanze vorhanden sind. Zum Zeitpunkt der Identifikation von NIC1 war keines dieser Gene charakterisiert. Außer für das MATE-Protein ERC1 aus S. cerevisiae gab es keine Studien zu eukaryotischen Mitgliedern dieser großen Familie von Membranproteinen. Anhand NIC1 wurden heterologe Expressionssysteme zur funktionellen Charakterisierung von MATE-Proteinen aus Pflanzen etabliert. Die cDNA von NIC1 wurde nach ihrer Klonierung in X. laevis Oozyten und S. cerevisiae exprimiert. In S. cerevisiae erhöhte die NIC1-Expression die Lithumtoleranz der Hefen und führte zu einer Verminderung der Natriumtoleranz. Parallele Versuche mit NIC2 und NIC4 (in den Diplomarbeiten von Blazej Dolniak und Mandy Kursawe) zeigten, dass auch diese beiden Proteine die Salztoleranz von S. cerevisiae beeinflussten. Während NIC2 die Lithium- und Natriumtoleranz erhöhte, führte NIC4-Expresion zu einer höheren Sensibilität gegenüber diesen beiden Kationen. Die unterschiedlichen Eigenschaften der drei homologen Proteine zeigten sich auch bei ihrer Expression in X. laevis Oozyten. NIC1 induzierte in den Oozyten auswärts gerichtete Chloridströme, die spannungsabhängig waren und durch mikromolare Konzentrationen der trivalenten Kationen Lanthan oder Gadolinium inhibiert werden konnten. NIC4 induzierte Barium-inhibierbare Kaliumströme, die spannungsunabhängig waren. Für NIC2 ließ sich in diesem Expressionssystem keine Aktivität detektieren. Zur Untersuchung der NIC1-Funktion in planta wurde die Genaktivität in transgenen Pflanzen lokalisiert und reduziert. Die NIC1-Genexpression war hauptsächlich in den vaskulären Geweben der Pflanze detektierbar und einer Verminderung des NIC1-Transkriptgehalts beeinflusste die Entwicklungsgeschwindigkeit der Pflanzen. Sie entwickelten sich deutlich langsamer als der parallel kultivierte Wildtyp. Der deutliche Phänotyp bei Veränderung der Genaktivität von nur einem der mindestens 56 vorhandenen MATE-Gene in A. thaliana zeigte, dass vermutlich keine weitere MATE-Isoform in der Lage ist, die Funktion von NIC1 zu übernehmen.