TY - JOUR A1 - Batram, Christopher A1 - Jones, Nivola G. A1 - Janzen, Christian J. A1 - Markert, Sebastian M. A1 - Engstler, Markus T1 - Expression site attenuation mechanistically links antigenic variation and development in Trypanosoma brucei JF - eLife N2 - We have discovered a new mechanism of monoallelic gene expression that links antigenic variation, cell cycle, and development in the model parasite Trypanosoma brucei. African trypanosomes possess hundreds of variant surface glycoprotein (VSG) genes, but only one is expressed from a telomeric expression site (ES) at any given time. We found that the expression of a second VSG alone is sufficient to silence the active VSG gene and directionally attenuate the ES by disruptor of telomeric silencing-1B (DOT1B)-mediated histone methylation. Three conserved expression-site-associated genes (ESAGs) appear to serve as signal for ES attenuation. Their depletion causes G1-phase dormancy and reversible initiation of the slender-to-stumpy differentiation pathway. ES-attenuated slender bloodstream trypanosomes gain full developmental competence for transformation to the tsetse fly stage. This surprising connection between antigenic variation and developmental progression provides an unexpected point of attack against the deadly sleeping sickness. KW - antigenic variation KW - expression site attenuation KW - developmental reprogramming KW - cell biology KW - genes and chromosomes KW - Trypanosoma brucei KW - variant surface glycoprotein (VSG) KW - monoallelic expression Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-119727 SN - 2050-084X VL - 3 IS - e02324 ER - TY - JOUR A1 - Baur, Stefanie A1 - Rautenberg, Maren A1 - Faulstich, Manuela A1 - Grau, Timo A1 - Severin, Yannik A1 - Unger, Clemens A1 - Hoffmann, Wolfgang H. A1 - Rudel, Thomas A1 - Autenrieth, Ingo B. A1 - Weidenmaier, Christopher T1 - A Nasal Epithelial Receptor for Staphylococcus aureus WTA Governs Adhesion to Epithelial Cells and Modulates Nasal Colonization JF - PLOS PATHOGENS N2 - Nasal colonization is a major risk factor for S. aureus infections. The mechanisms responsible for colonization are still not well understood and involve several factors on the host and the bacterial side. One key factor is the cell wall teichoic acid (WTA) of S. aureus, which governs direct interactions with nasal epithelial surfaces. We report here the first receptor for the cell wall glycopolymer WTA on nasal epithelial cells. In several assay systems this type F-scavenger receptor, termed SREC-I, bound WTA in a charge dependent manner and mediated adhesion to nasal epithelial cells in vitro. The impact of WTA and SREC-I interaction on epithelial adhesion was especially pronounced under shear stress, which resembles the conditions found in the nasal cavity. Most importantly, we demonstrate here a key role of the WTA-receptor interaction in a cotton rat model of nasal colonization. When we inhibited WTA mediated adhesion with a SREC-I antibody, nasal colonization in the animal model was strongly reduced at the early onset of colonization. More importantly, colonization stayed low over an extended period of 6 days. Therefore we propose targeting of this glycopolymer-receptor interaction as a novel strategy to prevent or control S. aureus nasal colonization. KW - SREC-I KW - clumping factor-B KW - scavender receptor KW - teichoic acids KW - surface proteins KW - cotton rats KW - carriage KW - determinant KW - infections KW - expression Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-116280 SN - 1553-7374 VL - 10 IS - 5 ER - TY - JOUR A1 - Bensaad, Karim A1 - Favaro, Elena A1 - Lewis, Caroline A. A1 - Peck, Barrie A1 - Lord, Simon A1 - Collins, Jennifer M. A1 - Pinnick, Katherine E. A1 - Wigfield, Simon A1 - Buffa, Francesca M. A1 - Li, Ji-Liang A1 - Zhang, Qifeng A1 - Wakelam, Michael J. O. A1 - Karpe, Fredrik A1 - Schulze, Almut A1 - Harris, Adrian L. T1 - Fatty Acid Uptake and Lipid Storage Induced by HIF-1 alpha Contribute to Cell Growth and Survival after Hypoxia-Reoxygenation JF - Cell Reports N2 - An in vivo model of antiangiogenic therapy allowed us to identify genes upregulated by bevacizumab treatment, including Fatty Acid Binding Protein 3 (FABP3) and FABP7, both of which are involved in fatty acid uptake. In vitro, both were induced by hypoxia in a hypoxia-inducible factor-1 alpha (HIF-1 alpha)-dependent manner. There was a significant lipid droplet (LD) accumulation in hypoxia that was time and O-2 concentration dependent. Knockdown of endogenous expression of FABP3, FABP7, or Adipophilin (an essential LD structural component) significantly impaired LD formation under hypoxia. We showed that LD accumulation is due to FABP3/7-dependent fatty acid uptake while de novo fatty acid synthesis is repressed in hypoxia. We also showed that ATP production occurs via beta-oxidation or glycogen degradation in a cell-type-dependent manner in hypoxia-reoxygenation. Finally, inhibition of lipid storage reduced protection against reactive oxygen species toxicity, decreased the survival of cells subjected to hypoxia-reoxygenation in vitro, and strongly impaired tumorigenesis in vivo. KW - inducible factor-I KW - binding protein KW - triglyceride accumulation KW - cancer cell KW - complex-III KW - beta-oxidation KW - metabolism KW - lipogenesis KW - proliferation KW - resistance Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-115162 SN - 2211-1247 VL - 9 IS - 1 ER - TY - JOUR A1 - Benz, Roland A1 - Maier, Elke A1 - Bauer, Susanne A1 - Ludwig, Albrecht T1 - The Deletion of Several Amino Acid Stretches of Escherichia coli Alpha-Hemolysin (HlyA) Suggests That the Channel-Forming Domain Contains Beta-Strands JF - PLOS ONE N2 - Escherichia coli α-hemolysin (HlyA) is a pore-forming protein of 110 kDa belonging to the family of RTX toxins. A hydrophobic region between the amino acid residues 238 and 410 in the N-terminal half of HlyA has previously been suggested to form hydrophobic and/or amphipathic α-helices and has been shown to be important for hemolytic activity and pore formation in biological and artificial membranes. The structure of the HlyA transmembrane channel is, however, largely unknown. For further investigation of the channel structure, we deleted in HlyA different stretches of amino acids that could form amphipathic β-strands according to secondary structure predictions (residues 71–110, 158–167, 180–203, and 264–286). These deletions resulted in HlyA mutants with strongly reduced hemolytic activity. Lipid bilayer measurements demonstrated that HlyAΔ71–110 and HlyAΔ264–286 formed channels with much smaller single-channel conductance than wildtype HlyA, whereas their channel-forming activity was virtually as high as that of the wildtype toxin. HlyAΔ158–167 and HlyAΔ180–203 were unable to form defined channels in lipid bilayers. Calculations based on the single-channel data indicated that the channels generated by HlyAΔ71–110 and HlyAΔ264–286 had a smaller size (diameter about 1.4 to 1.8 nm) than wildtype HlyA channels (diameter about 2.0 to 2.6 nm), suggesting that in these mutants part of the channel-forming domain was removed. Osmotic protection experiments with erythrocytes confirmed that HlyA, HlyAΔ71–110, and HlyAΔ264–286 form defined transmembrane pores and suggested channel diameters that largely agreed with those estimated from the single-channel data. Taken together, these results suggest that the channel-forming domain of HlyA might contain β-strands, possibly in addition to α-helical structures. KW - membrane potential KW - molecular mass KW - cations KW - membrane structures KW - membrane proteins KW - lipid bilayer KW - red blood cells KW - toxins Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-118115 SN - 1932-6203 VL - 9 IS - 12 ER - TY - THES A1 - Bettaga, Noomen T1 - Bedeutung der NO-sensitiven Guanylyl Cyclase bei der Angiogenese und der Arteriogenese in der Maus T1 - Role of NO-sensitive guanylyl cyclase in angiogenesis and arteriogenesis in mice N2 - Stickstoffmonoxid (NO) spielt eine wichtige Rolle bei Gefäßremodelling-Prozessen wie Angiogenese und Arteriogenese. Die NO-Synthese im Gefäßsystem wird hauptsächlich durch die endotheliale NO-Synthase (eNOS) gewährleistet. Sie kann durch verschiedene Faktoren wie Scherkräfte und Zytokine wie der vaskuläre endotheliale Wachstumsfaktor (VEGF) reguliert werden. VEGF ist ein wichtiger Stimulator der Angiogenese und wird während dieses Prozesses hochreguliert. Die meisten physiologischen Effekte von NO werden durch die NO-sensitive Guanylyl-Cyclase (NO-GC) vermittelt. Als Hauptrezeptor für NO produziert die NO-GC den sekundären Botenstoff cyklisches Guanosinmonophosphat (cGMP) und führt dadurch zur Stimulation der verschiedenen Effektoren wie z.B. der PKG. Ob die Wirkung von NO in Angiogenese und Arteriogenese ebenfalls durch NO-GC vermittelt wird, war bis zum Beginn dieser Arbeit noch unklar. Die NO-GC besteht aus zwei Untereinheiten (α und ß). Die Deletion der ß1-Untereinheit in Mäusen resultiert in einer vollständigen Knockout Maus (GCKO). Mithilfe des Cre-LoxP-Systems wurden zusätzlich zellspezifische Knockout-Mäuse für glatte Muskelzellen (SMC-GCKO) und Endothelzellen (EC-GCKO) generiert. Um die Rolle der NO-GC in der Angiogenese und Arteriogenese zu untersuchen, wurden drei gut etablierte Methoden benutzt. Im ersten Teil des Projekts sollte die Expression der NO-GC in Endothelzellen untersucht werden. Zu diesem Zweck wurde die reverse Transkriptase-Polymerase-Kettenreaktion (RT-PCR) benutzt. Die Ergebnisse zeigen, dass die NO-GC in Endothelzellen der Lunge nur äußerst gering wenig exprimiert ist. Durch den Aortenring-Assay wurde eine Rolle der NO-GC bei der VEGF-vermittelten Angiogenese festgestellt. Dabei zeigte sich eine stärkere Angiogeneserate bei globaler Abwesenheit der NO-GC. Bei Fehlen der NO-GC ausschließlich in Endothelzellen zeigte sich kein Unterschied in den aussprossenden Aorten im Vergleich zu den Kontroll-Tieren. Dies zeigt, dass die NO-GC in Endothelzellen sehr wahrscheinlich keine Rolle bei der VEGF-vermittelten Angiogenese spielt. Im zweiten Teil wurde die Rolle der NO-GC bei der Angiogenese in einem in vivo-Modell untersucht. In dem Modell der Sauerstoff-induzierten-Retinopathie zeigten die GCKO-Mäuse eine verringerte Vaso-Obliteration, eine verlangsamte Angiogenese und eine erhöhte Tuft-Bildung. Ähnliche Ergebnisse wurden bei den SMC-GCKO-Tieren beobachtet. EC-GCKO-Mäuse zeigten eine gegenüber den Kontroll-Tieren unveränderte Vaso-Obliteration, Angiogeneserate und Tuft-Bildung. Diese Ergebnisse lassen darauf schließen, dass die NO-GC in Endothelzellen keine Rolle spielt. Immunfluoreszenz-Aufnahmen zeigten die Expression von NO-GC in Perizyten der Gefäßkapillaren der Mausretina. Daher könnte die NO-GC in diesem Zelltyp letztendlich für die Effekte bei den GCKO- und SMC-GCKO-Tieren verantwortlich sein. Im letzten Teil dieser Arbeit wurde eine Versuchsreihe unter Anwendung des Hinterlauf-Ischämie-Modells durchgeführt. Hierbei entwickelten die Pfoten aller GCKO- und teilweise der SMC-GCKO-Tiere nach der Ligation der Femoralarterie eine Nekrose. Die Regeneration der Hinterläufe der EC-GCKO-Tiere nach der Operation verlief normal. Diese Ergebnisse schließen eine bedeutende Rolle der NO-GC in Endothelzellen aus, zeigen allerdings, dass die NO-GC in den glatten Muskelzellen essentiell für den Arteriogenese-Prozess ist. Zusammengefasst führt die Deletion der NO-GC in glatten Muskelzellen und wahrscheinlich auch in Perizyten zur einer verlangsamten Angiogenese und Inhibierung der Arteriogenese. N2 - Nitric oxide (NO) plays an important role in vascular remodelling processes such as angiogenesis and arteriogenesis. The synthesis of NO in the vascular system is ensured mainly by endothelial NO synthase (eNOS). It can be regulated by a number of factors, such as shear stress and cytokines like the vascular endothelial growth factor (VEGF). VEGF is an important stimulator of angiogenesis and is upregulated during this process. Most of the physiological effects of NO are mediated by the NO-sensitive guanylyl cyclase (NO-GC). As the main receptor for NO, NO-GC produces the second messenger cyclic guanosine monophosphate (cGMP) and thereby leads to a variety of physiological effects. However, whether the effects of NO in angiogenesis and arteriogenesis are also mediated by NO-GC is still unclear. NO-GC consists of two subunits (α and ß). The deletion of the ß1 subunit in mice results in a global knockout mouse (GCKO). Using the Cre-LoxP system we also generated smooth muscle cell-specific (SMC GCKO) and endothelial cell-specific knockout mice (EC GCKO). To investigate the role of NO-GC in angiogenesis and arteriogenesis, three well-established methods have been used. In the first part of the project, the expression of the NO-GC in endothelial cells should be investigated. By using the reverse transcription polymerase chain reaction method (RT-PCR), the results show a very weak expression of the NO-GC in endothelial cells of the lung. A role for NO-GC in the VEGF-mediated angiogenesis was ascertained by the aortic ring assay. The results show an increased angiogenesis in the global absence of NO-GC. However, the EC-GCKO shows no difference compared to control mice. This indicates that NO-GC in endothelial cells is unlikely to play a major role in VEGF-mediated angiogenesis. In the second part of the project, the role of NO-GC in angiogenesis was investigated in an in vivo model. In the oxygen induced-retinopathy model (OIR), GCKO mice showed reduced vaso-obliteration, slowed angiogenesis and increased tuft formation. Similar results were observed in the SMC-GCKO animals. In contrast, EC-GCKO mice showed vaso-obliteration as well as angiogenesis rate and tuft formation similar to those seen in control animals. The results of this experiment suggest that NO-GC in endothelial cells is not involved in vaso-obliteration, physiological angiogenesis and tuft formation. Immunhistochemical analyses showed the expression of NO-GC in pericytes of the vascular capillaries of the mouse retina. Therefore, NO-GC in this cell type could be responsible for the effects in GCKO- and SMC-GCKO animals. In the last part of this thesis, hindlimb-ischemia experiments were performed. For this purpose, the paws of all GCKO- and some SMC-GCKO animals showed necrosis after ligation of the femoral artery. The regeneration of legs from EC-GCKO animals after the operation was normal. These results exclude a major role of NO-GC in endothelial cells, but show that NO-GC in smooth muscle cells is essential in the arteriogenesis process. In summary, the deletion of NO-GC in smooth muscle cells and probably also in pericytes leads to a slowed angiogenesis and inhibits arteriogenesis. KW - Guanylylcyclase KW - cGMP KW - Stickstoffmonoxid KW - Angiogenese KW - Arteriogenese KW - Maus Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-111284 ER - TY - JOUR A1 - Breeze, Tom D. A1 - Vaissiere, Bernhard E. A1 - Bommarco, Riccardo A1 - Petanidou, Theodora A1 - Seraphides, Nicos A1 - Kozak, Lajos A1 - Scheper, Jeroen A1 - Biesmeijer, Jacobus C. A1 - Kleijn, David A1 - Gyldenkærne, Steen A1 - Moretti, Marco A1 - Holzschuh, Andrea A1 - Steffan-Dewenter, Ingolf A1 - Stout, Jane C. A1 - Pärtel, Meelis A1 - Zobel, Martin A1 - Potts, Simon G. T1 - Agricultural Policies Exacerbate Honeybee Pollination Service Supply-Demand Mismatches Across Europe JF - PLOS ONE N2 - Declines in insect pollinators across Europe have raised concerns about the supply of pollination services to agriculture. Simultaneously, EU agricultural and biofuel policies have encouraged substantial growth in the cultivated area of insect pollinated crops across the continent. Using data from 41 European countries, this study demonstrates that the recommended number of honeybees required to provide crop pollination across Europe has risen 4.9 times as fast as honeybee stocks between 2005 and 2010. Consequently, honeybee stocks were insufficient to supply >90% of demands in 22 countries studied. These findings raise concerns about the capacity of many countries to cope with major losses of wild pollinators and highlight numerous critical gaps in current understanding of pollination service supplies and demands, pointing to a pressing need for further research into this issue. KW - economy services KW - fruit set KW - sequential introduction KW - enhance KW - biodiversity KW - abundance KW - declines KW - crops KW - colonies KW - density Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-117692 SN - 1932-6203 VL - 9 IS - 1 ER - TY - JOUR A1 - Brehm, Klaus A1 - Hemer, Sarah A1 - Konrad, Christian A1 - Spiliotis, Markus A1 - Koziol, Uriel A1 - Schaack, Dominik A1 - Förster, Sabine A1 - Gelmedin, Verena A1 - Stadelmann, Britta A1 - Dandekar, Thomas A1 - Hemphill, Andrew T1 - Host insulin stimulates Echinococcus multilocularis insulin signalling pathways and larval development N2 - Background The metacestode of the tapeworm Echinococcus multilocularis is the causative agent of alveolar echinococcosis, a lethal zoonosis. Infections are initiated through establishment of parasite larvae within the intermediate host’s liver, where high concentrations of insulin are present, followed by tumour-like growth of the metacestode in host organs. The molecular mechanisms determining the organ tropism of E. multilocularis or the influences of host hormones on parasite proliferation are poorly understood. Results Using in vitro cultivation systems for parasite larvae we show that physiological concentrations (10 nM) of human insulin significantly stimulate the formation of metacestode larvae from parasite stem cells and promote asexual growth of the metacestode. Addition of human insulin to parasite larvae led to increased glucose uptake and enhanced phosphorylation of Echinococcus insulin signalling components, including an insulin receptor-like kinase, EmIR1, for which we demonstrate predominant expression in the parasite’s glycogen storage cells. We also characterized a second insulin receptor family member, EmIR2, and demonstrated interaction of its ligand binding domain with human insulin in the yeast two-hybrid system. Addition of an insulin receptor inhibitor resulted in metacestode killing, prevented metacestode development from parasite stem cells, and impaired the activation of insulin signalling pathways through host insulin. Conclusions Our data indicate that host insulin acts as a stimulant for parasite development within the host liver and that E. multilocularis senses the host hormone through an evolutionarily conserved insulin signalling pathway. Hormonal host-parasite cross-communication, facilitated by the relatively close phylogenetic relationship between E. multilocularis and its mammalian hosts, thus appears to be important in the pathology of alveolar echinococcosis. This contributes to a closer understanding of organ tropism and parasite persistence in larval cestode infections. Furthermore, our data show that Echinococcus insulin signalling pathways are promising targets for the development of novel drugs. KW - Cestode KW - Tapeworm KW - Echinococcus KW - Echinococcosis KW - Insulin KW - Receptor kinase KW - Kinase inhibitor KW - Host-parasite interaction Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-110357 ER - TY - JOUR A1 - Brehm, Klaus A1 - Koziol, Uriel A1 - Rauschendorfer, Theresa A1 - Rodríguez, Luis Zanon A1 - Krohne, Georg T1 - The unique stem cell system of the immortal larva of the human parasite Echinococcus multilocularis N2 - Background It is believed that in tapeworms a separate population of undifferentiated cells, the germinative cells, is the only source of cell proliferation throughout the life cycle (similar to the neoblasts of free living flatworms). In Echinococcus multilocularis, the metacestode larval stage has a unique development, growing continuously like a mass of vesicles that infiltrate the tissues of the intermediate host, generating multiple protoscoleces by asexual budding. This unique proliferation potential indicates the existence of stem cells that are totipotent and have the ability for extensive self-renewal. Results We show that only the germinative cells proliferate in the larval vesicles and in primary cell cultures that undergo complete vesicle regeneration, by using a combination of morphological criteria and by developing molecular markers of differentiated cell types. The germinative cells are homogeneous in morphology but heterogeneous at the molecular level, since only sub-populations express homologs of the post-transcriptional regulators nanos and argonaute. Important differences are observed between the expression patterns of selected neoblast marker genes of other flatworms and the E. multilocularis germinative cells, including widespread expression in E. multilocularis of some genes that are neoblast-specific in planarians. Hydroxyurea treatment results in the depletion of germinative cells in larval vesicles, and after recovery following hydroxyurea treatment, surviving proliferating cells grow as patches that suggest extensive self-renewal potential for individual germinative cells. Conclusions In E. multilocularis metacestodes, the germinative cells are the only proliferating cells, presumably driving the continuous growth of the larval vesicles. However, the existence of sub-populations of the germinative cells is strongly supported by our data. Although the germinative cells are very similar to the neoblasts of other flatworms in function and in undifferentiated morphology, their unique gene expression pattern and the evolutionary loss of conserved stem cells regulators suggest that important differences in their physiology exist, which could be related to the unique biology of E. multilocularis larvae. KW - Cestoda KW - Echinococcus KW - Neoblast KW - Germinative cell KW - Stem cell KW - Nanos KW - Argonaute KW - Mucin KW - Alkaline phosphatase Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-110315 ER - TY - JOUR A1 - Chen, Yi-chun A1 - Gerber, Bertram T1 - Generalization and discrimination tasks yield concordant measures of perceived distance between odours and their binary mixtures in larval Drosophila JF - The Journal of Experimental Biology N2 - Similarity between odours is notoriously difficult to measure. Widely used behavioural approaches in insect olfaction research are cross-adaptation, masking, as well as associative tasks based on olfactory learning and the subsequent testing for how specific the established memory is. A concern with such memory-based approaches is that the learning process required to establish an odour memory may alter the way the odour is processed, such that measures of perception taken at the test are distorted. The present study was therefore designed to see whether behavioural judgements of perceptual distance are different for two different memory-based tasks, namely generalization and discrimination. We used odour-reward learning in larval Drosophila as a study case. In order to challenge the larvae's olfactory system, we chose to work with binary mixtures and their elements (1-octanol, n-amyl acetate, 3-octanol, benzaldehyde and hexyl acetate). We determined the perceptual distance between each mixture and its elements, first in a generalization task, and then in a discrimination task. It turns out that scores of perceptual distance are correlated between both tasks. A re-analysis of published studies looking at element-to-element perceptual distances in larval reward learning and in adult punishment learning confirms this result. We therefore suggest that across a given set of olfactory stimuli, associative training does not grossly alter the pattern of perceptual distances. KW - discrimination KW - drosophila melanogaster KW - generalization KW - memory KW - olfaction KW - perception Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-121625 VL - 217 IS - 12 ER - TY - THES A1 - Claßen, Alice T1 - Diversity, traits and ecosystem services of pollinators along climate and land use gradients on Mount Kilimanjaro T1 - Diversität, Merkmale und Ökosystemfunktionen von Bestäubern entlang von Klima- und Landnutzungsgradienten am Kilimandscharo N2 - Since more than two centuries naturalists are fascinated by the profound changes in biodiversity observed along climatic gradients. Although the theories explaining changes in the diversity and the shape of organisms along climatic gradients belong to the foundations of modern ecology, our picture on the spatial patterns and drivers of biodiversity is far from being complete. Ambiguities in theory and data are common and past work has been strongly concentrated on plants and vertebrates. In the last two decades, interest in the fundamental processes structuring diversity along climatic gradients gained new impetus as they are expected to improve our understanding about how ecosystems will respond to global environmental changes. Global temperatures are rising faster than ever before; natural habitats are transformed into agricultural land and existing land use systems get more and more intensified to meet the demands of growing human populations. The fundamental shifts in the abiotic and biotic environment are proclaimed to affect ecosystems all over the world; however, precise predictions about how ecosystems respond to global changes are still lacking. We investigated diversity, traits and ecosystem services of wild bees along climate and land use gradients on Mount Kilimanjaro (Tanzania, East Africa). Wild bees play a major role in ecosystems, as they contribute to the reproduction and performance of wild and crop plants. Their responsiveness to environmental changes is therefore of high ecological and economic importance. Temperature and energy resources have often been suggested to be the main determinants of global and local species richness, but the mechanisms behind remain poorly understood. In the study described in chapter II we analyzed species richness patterns of wild bees along climate and land use gradients on Mount Kilimanjaro and disentangled the factors explaining most of the changes in bee richness. We found that floral resources had a weak but significant effect on pollinator abundance, which in turn was positively related to species richness. However, temperature was the strongest predictor of species richness, affecting species richness both directly and indirectly by positively influencing bee abundances. We observed higher levels of bee-flower-interactions at higher temperatures, independently of flower and bee abundances. This suggests that temperature restricts species richness by constraining the exploitation of resources by ectotherms. Current land use did not negatively affect species richness. We conclude that the richness of bees is explained by both temperature and resource availability, whereas temperature plays the dominant role as it limits the access of ectotherms to floral resources and may accelerate ecological and evolutionary processes that drive the maintenance and origination of diversity. Not only species numbers, but also morphological traits like body size are expected to be shaped by both physiological and energetic constraints along elevational gradients. Paradoxically, Bergmann´s rule predicts increases of body sizes in cooler climates resulting from physiological constraints, while species-energy theory suggests declines in the mean body size of species caused by increased extinction probabilities for large-bodied species in low-energy habitats. In chapter III we confronted this ambiguity with field data by studying community-wide body size variation of wild bees on Mt. Kilimanjaro. We found that along a 3680 m elevational gradient bee individuals became on average larger within species, while large species were increasingly absent from high-elevational communities. This demonstrates, on the one hand, how well-established, but apparently contrasting ecological theories can be merged through the parallel consideration of different levels of biological organization. On the other hand it signals that the extinction risk in the course of environmental change is not equally distributed among species within a community. Land use intensification is known to threaten biodiversity, but the consequences for ecosystem services are still a matter of debate. In chapter IV, we experimentally tested the single and combined contributions of pest predators and pollinators to coffee production along a land use intensification gradient on Mount Kilimanjaro. We found that pest predation increased fruit set by on average 9%, while pollination increased fruit weight of coffee by on average 7.4%. Land use had no significant effect on both ecosystem services. However, we found that in coffee plantations with most intensified land use, pollination services were virtually exclusively provided by the honey bee (Apis mellifera). The reliance on a single pollinator species is risky, as possible declines of that species may directly lower pollination services, resulting in yield losses. In contrast, pollination services in structurally complex homegardens were found to be provided by a diverse pollinator community, increasing the stability of pollination services in a long term. We showed that on Mount Kilimanjaro pollinator communities changed along elevational gradients in terms of species richness (chapter II) and trait composition (chapter III). Temperature and the temperature-mediated accessibility of resources were identified as important predictors of these patterns, which contributes to our fundamental understanding about the factors that shape ectothermic insect communities along climatic gradients. The strong temperature-dependence of pollinators suggests that temperature shifts in the course of global change are likely to affect pollinator communities. Pollinators might either profit from rising temperatures, or shift to higher elevations, which could result in related biotic attrition in the lowland with consequences for the provision of ecosystem services in cropping systems. Up to now, land use intensification had no significant impact on the diversity of pollinator communities and their ecosystem services. Pollinators might profit from the strong landscape heterogeneity in the region and from the amount of flower resources in the understory of cropping systems. However,progressing homogenization of the landscape and the pronounced application of pesticides could result in reduced diversity and dominance of single species, as we already found in sun coffee plantations. Such shifts in community compositions could threaten the stability of ecosystem services within cropping and natural systems in a long term. N2 - Die Biodiversität auf der Erde ist nicht gleichmäßig verteilt, sondern verändert sich entlang klimatischer Gradienten – ein Phänomen, das Naturwissenschaftler schon seit mehr als zwei Jahrhunderten fasziniert. Über die Mechanismen, welche die Verteilung von Arten entlang von Klimazonen bestimmen, besteht nach wie vor keine Einigkeit, auch wenn viele der hier hervorgebrachten Theorien zu den Grundlagen der modernen Ökologie gehören. Ambivalenzen in den Erklärungsmodellen und erhobenen Daten sind häufig und bisherige Studien konzentrierten sich vorrangig auf Pflanzen und Vertebraten, während andere Taxa weniger Beachtung fanden. Die Unsicherheit über die Auswirkungen des globalen Wandels auf Ökosysteme und die Konsequenzen für Ökosystemdienstleistungen setzte neue Impulse im Bereich der Biodiversitätsforschung. Temperaturen steigen so schnell wie nie zuvor; natürliche Habitate werden zunehmend in Agrarflächen umgewandelt und bestehende Landwirtschaftssysteme werden intensiviert. Präzise Vorhersagen darüber, wie Ökosysteme auf solch drastische Umweltveränderungen reagieren, fehlen jedoch weitgehend. In dieser Dissertation wird gezeigt, wie sich die Artenvielfalt, morphologische Merkmale und Ökosystemfunktionen von Bienen entlang von Klima- und Landnutzungsgradienten des Kilimandscharos (Tansania, Ostafrika) verändern. Bienen spielen eine wichtige Rolle in Ökosystemen, da sie als Bestäuber zur Reproduktion und Produktivität von Wild- und Nutzpflanzen beitragen. Die Veränderung ihres Artenreichtums, ihrer Merkmale und ihrer Ökosystemdienstleistungen entlang von Umweltgradienten ist somit von großer ökologischer und ökonomischer Relevanz. Temperatur und die Verfügbarkeit von Ressourcen sind die Faktoren, mit denen globale und lokale Muster von Diversität am häufigsten erklärt werden. Die kausalen Zusammenhänge über welche die Temperatur und die Verfügbarkeit von Ressourcen eine Erhöhung der Artenvielfalt bewirken, sind jedoch nach wie vor unklar. Im zweiten Kapitel dieser Arbeit wurde untersucht, wie sich der Artenreichtum von Bienen entlang von Klima- und Landnutzungsgradienten des Kilimandscharos verändert und welche Faktoren für diese Veränderungen verantwortlich sind. Blühressourcen hatten einen schwachen, aber signifikanten Einfluss auf die Bestäuberabundanzen, welche wiederum einen Großteil des Artenreichtums erklärten. Insgesamt hatte die Temperatur jedoch einen deutlich stärken Einfluss auf die Artenvielfalt als die Verfügbarkeit von Blühressourcen: Die Temperatur wirkte sich direkt, möglicherweise über einen Erhöhung von Speziationsraten, und indirekt, über eine Erhöhung der Bienenabundanz, auf die Artenvielfalt von Bienen aus. Zusätzlich konnte beobachtet werden, dass die Blütenbesuche von Bienen unabhängig von der Blütendichte und der Bienenabundanz, mit einem Anstieg der Temperatur zunahmen. Aus diesen Beobachtungen folgern wir, dass bei Ektothermen die Nutzbarkeit von Ressourcen durch die Temperatur gesteuert sein könnte. Die Untersuchung morphologischer Merkmale entlang von Umweltgradienten erlaubt es, deterministische von stochastischen Prozessen bei der Zusammensetzung von Artengemeinschaften zu unterscheiden. Während bei stochastischen Prozessen Merkmale entlang von Umweltgradienten zufällig aus Artengemeinschaften ausscheiden sollten, wird im Falle deterministischer Prozesse ein gerichtetes Muster erwartet. Unter den deterministischen Prozessen konkurrieren bezüglich der Körpergröße zwei scheinbar konträre Theorien: Während die Bergmannsche Regel vorhersagt, dass große Tiere, aufgrund eines verbesserten Oberflächen-Volumen-Verhältnisses, einen Vorteil in kühlen Regionen haben, weist die Arten-Energie-Theorie größeren Arten eine erhöhte Aussterbewahrscheinlichkeit in energielimitierten, kühlen Gebieten zu, so dass die mittlere Körpergröße von Lebensgemeinschaften bei kälterer Temperatur sinken sollte. Im dritten Kapitel dieser Dissertation untersuchten wir, ob sich morphologische Merkmale von Wildbienen mit zunehmender Höhe verändern. Dabei betrachteten wir nicht nur Merkmalsveränderungen innerhalb von Artengemeinschaften, sondern auch innerartliche Veränderungen. Sowohl physiologische als auch energetische Restriktionen prägten die Merkmalskompositionen, allerdings auf unterschiedlichen biologischen Ebenen. So nahm die Körpergröße innerhalb von Arten mit der Höhe im Durchschnitt zu (=Bergmannsche Regel), während auf Gemeinschaftsebene kleinere Arten die Hochgebirgsregionen dominierten (=energetische Restriktion). Die parallele Betrachtung der intra- und interspezifischen Ebene ermöglichte es uns, scheinbar konträre ökologische Theorien zusammenzuführen. Zudem konnten wir zeigen, dass Merkmale nicht zufällig, sondern gerichtet aus Artengemeinschaften gefiltert werden. Landnutzungsintensivierung bedroht Biodiversität, aber die Konsequenzen für Ökosystemdienstleistungen sind nach wie vor ungewiss. Im vierten Kapitel dieser Arbeit prüften wir mit Hilfe von einzelnen und kombinierten Bestäuber- und Prädatorausschlussexperimenten, welchen Beitrag Bestäuber, Vögel und Fledermäuse in verschiedenen Anbausystemen zur Kaffeeproduktion am Kilimandscharo leisten. Wir zeigten, dass sich Bestäuber und Prädatoren in ihren Effekten ergänzten: Während Bestäuber eine Steigerung des Kaffeebohnengewichtes um durchschnittlich 7.4% bewirkten, konnte durch die Prädation von Schädlingen der Fruchtansatz des Kaffees um durchschnittlich 9% gesteigert werden. Landwirtschaftliche Intensivierung, von komplexen Waldwirtschaftssystemen, über beschattete Kaffeeplantagen, bis hin zu Sonnenplantagen hatte keinen negativen Effekt auf die Ökosystemdienstleistungen von Bestäubern und Prädatoren. Wir konnten jedoch nachweisen, dass in Waldwirtschaftssystemen eine diverse Bestäubergemeinschaft den Kaffee bestäubt, während in Sonnenplantagen fast ausschließlich die Honigbiene als Bestäuber fungiert. Eine solche Verschiebung der Bestäuber-komposition könnte die langfristige Stabilität intensiv genutzter Flächen gefährden. In dieser Dissertation zeigten wir, wie sich Bestäubergemeinschaften am Kilimandscharo entlang von Höhengradienten bezüglich ihrer Artenvielfalt (Kapitel II) und ihrer Merkmale (Kapitel III) verändern. Temperatur und temperatur-gesteuerte Ressourcennutzbarkeit wurden als maßgebende Determinanten dieser Muster identifiziert. Damit wurde ein Beitrag zur Identifikation von Gesetzmäßigkeiten in der Verteilung ektothermer Insekten entlang von Klimagradienten geleistet. Unsere Ergebnisse weisen darauf hin, dass Klimaveränderungen im Zuge des globalen Wandels Konsequenzen für Bestäubergemeinschaften haben könnten. Eventuell könnten Bestäuber von den steigenden Temperaturen profitieren. Gleichsam könnte es aber auch zu einer Verschiebung von Bestäubern in höher gelegene Regionen und zu einem daran gekoppelten Einbruch der Bestäubungsleistungen in tiefliegenden Kulturlandschaften kommen. Im Hinblick auf die Konsequenzen anthropogener Landnutzung wurde festgestellt, dass die landwirtschaftliche Intensivierung am Kilimandscharo bisher keinen messbaren negativen Effekt auf die Ökosystemdienstleitungen von Bestäubern hatte. Die Bestäuber profitieren vermutlich von der starken Landschaftsheterogenität der Region und zahlreichen krautigen Blühressourcen im Unterwuchs von Agrarflächen. Eine zunehmende Homogenisierung der Landschaft und ein verstärkter Einsatz von Pestiziden könnten jedoch, wie auf Sonnenplantagen bereits zu finden, zu einer Dominanz von einigen wenigen Arten führen, welches zusammen mit der klimabedingten Artenverschiebung die langfristige Stabilität von Agrarsystemen und natürlichen Systemen gefährden könnte. KW - Kilimandscharo KW - Bestäuber KW - Biodiversität KW - Höhengradient KW - Landnutzungsgradient KW - Arten-Energy-Theory KW - species-energy-theory KW - elevational gradient KW - Kilimanjaro KW - pollinators KW - diversity KW - ecosystem service KW - ecology Y1 - 2014 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:20-opus-101292 ER -