@phdthesis{Schilder1999, author = {Schilder, Klaus}, title = {Safer without Sex?}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-1977}, school = {Universit{\"a}t W{\"u}rzburg}, year = {1999}, abstract = {Highly eusocial insect societies, such as all known ants, are typically characterized by a reproductive division of labor between queens, who are inseminated and reproduce, and virgin workers, who engage in foraging, nest maintenance and brood care. In most species workers have little reproductive options left: They usually produce haploid males by arrhenotokous parthenogenesis, both in the queenright and queenless condition. In the phylogenetically primitive subfamily Ponerinae reproductive caste dimorphism is much less pronounced: Ovarian morphology is rather similar in queens and workers, which additionally retain a spermatheca. In many ponerine species workers mate and may have completely replaced the queen caste. This similarity in reproductive potential provides for the evolution of diverse reproductive systems. In addition, it increases the opportunity for reproductive conflicts among nestmates substantially. Only in a handful of ant species, including Platythyrea punctata, workers are also able to rear diploid female offspring from unfertilized eggs by thelytokous parthenogenesis. The small ponerine ant P. punctata (Smith) is the only New World member of the genus reaching as far north as the southern USA, with its center of distribution in Central America and the West Indies. P. punctata occurs in a range of forest habitats including subtropical hardwood forests as well as tropical rain forests. In addition to queens, gamergates and thelytokous workers co-occur in the same species. This remarkable complexity of reproductive strategies makes P. punctata unique within ants and provides an ideal model system for the investigation of reproductive conflicts within the female caste. Colonies are usually found in rotten branches on the forest floor but may also be present in higher strata. Colonies contained on average 60 workers, with a maximum colony size of 148 workers. Queens were present in only ten percent of the colonies collected from Florida, but completely absent both from the populations studied in Barbados and Puerto Rico. Males were generally rare. In addition, morphological intermediates between workers and queens (so-called intercastes) were found in 16 colonies collected in Florida. Their thorax morphology varied from an almost worker-like to an almost queen-like thorax structure. Queen and intercaste size, however, did not differ from those of workers. Although workers taken from colonies directly after collection from the field engaged in aggressive interactions, nestmate discrimination ceased in the laboratory suggesting that recognition cues used are derived from the environment. Only one of six queens dissected was found to be inseminated but not fertile. Instead, in most queenless colonies, a single uninseminated worker monopolized reproduction by means of thelytokous parthenogenesis. A single mated, reproductive worker (gamergate) was found dominating reproduction in the presence of an inseminated alate queen only in one of the Florida colonies. The regulation of reproduction was closely examined in ten experimental groups of virgin laboratory-reared workers, in which one worker typically dominated reproduction by thelytoky despite the presence of several individuals with elongated, developing ovaries. In each group only one worker was observed to oviposit. Conflict over reproduction was intense consisting of ritualized physical aggression between some nestmates including antennal boxing, biting, dragging, leap and immobilization behaviors. The average frequency of interactions was low. Aggressive interactions allowed to construct non-linear matrices of social rank. On average, only five workers were responsible for 90 percent of total agonistic interactions. In 80 percent of the groups the rate of agonistic interactions increased after the experimental removal of the reproductive worker. While antennal boxing and biting were the most frequent forms of agonistic behaviors both before and after the removal, biting and dragging increased significantly after the removal indicating that agonistic interactions increased in intensity. Once a worker obtains a high social status it is maintained without the need for physical aggression. The replacement of reproductives by another worker did however not closely correlate with the new reproductive's prior social status. Age, however, had a profound influence on the individual rate of agonistic interactions that workers initiated. Especially younger adults (up to two month of age) and callows were responsible for the increase in observed aggression after the supersedure of the old reproductive. These individuals have a higher chance to become reproductive since older, foraging workers may not be able to develop their ovaries. Aggressions among older workers ceased with increasing age. Workers that already started to develop their ovaries should pose the greatest threat to any reproductive individual. Indeed, dissection of all experimental group revealed that aggression was significantly more often directed towards both individuals with undeveloped and developing ovaries as compared to workers that had degenerated ovaries. In all experimental groups reproductive dominance was achieved by callows or younger workers not older than four month. Age is a better predictor of reproductive dominance than social status as inferred from physical interactions. Since no overt conflict between genetical identical individuals is expected, in P. punctata the function of agonistic interactions in all-worker colonies, given the predominance of thelytokous parthenogenesis, remains unclear. Physical aggression could alternatively function to facilitate a smooth division of non-reproductive labor thereby increasing overall colony efficiency. Asexuality is often thought to constitute an evolutionary dead end as compared with sexual reproduction because genetic recombination is limited or nonexistent in parthenogenetic populations. Microsatellite markers were developed to investigate the consequences of thelytokous reproduction on the genetic structure of four natural populations of P. punctata. In the analysis of 314 workers taken from 51 colonies, low intraspecific levels of variation at all loci, expressed both as the number of alleles detected and heterozygosities observed, was detected. Surprisingly, there was almost no differentiation within populations. Populations rather had a clonal structure, with all individuals from all colonies usually sharing the same genotype. This low level of genotypic diversity reflects the predominance of thelytoky under natural conditions in four populations of P. punctata. In addition, the specificity of ten dinucleotide microsatellite loci developed for P. punctata was investigated in 29 ant species comprising four different subfamilies by cross-species amplification. Positive amplification was only obtained in a limited number of species indicating that sequences flanking the hypervariable region are often not sufficiently conserved to allow amplification, even within the same genus. The karyotype of P. punctata (2n = 84) is one of the highest chromosome numbers reported in ants so far. A first investigation did not show any indication of polyploidy, a phenomenon which has been reported to be associated with the occurrence of parthenogenesis. Thelytokous parthenogenesis does not appear to be a very common phenomenon in the Hymenoptera. It is patchily distributed and restricted to taxa at the distant tips of phylogenies. Within the Formicidae, thelytoky has been demonstrated only in four phylogenetically very distant species, including P. punctata. Despite its advantages, severe costs and constraints may have restricted its rapid evolution and persistence over time. The mechanisms of thelytokous parthenogenesis and its ecological correlates are reviewed for the known cases in the Hymenoptera. Investigating the occurrence of sexual reproduction in asexual lineages indicates that thelytokous parthenogenesis may not be irreversible. In P. punctata the occasional production of sexuals in some of the colonies may provide opportunity for outbreeding and genetic recombination. Thelytoky can thus function as a conditional reproductive strategy. Thelytoky in P. punctata possibly evolved as an adaptation to the risk of colony orphanage or the foundation of new colonies by fission. The current adaptive value of physical aggression and the production of sexuals in clonal populations, where relatedness asymmetries are virtually absent, however is less clear. Quite contrary, thelytoky could thereby serve as the stepping stone for the subsequent loss of the queen caste in P. punctata. Although P. punctata clearly fulfills all three conditions of eusociality, the evolution of thelytoky is interpreted as a first step in a secondary reverse social evolution towards a social system more primitive than eusociality.}, subject = {Ameisenstaat}, language = {en} } @phdthesis{Rueppell2000, author = {R{\"u}ppell, Olav}, title = {Queen size dimorphism in ants}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-1914}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2000}, abstract = {Many polymorphisms are linked to alternative reproductive strategies. In animals, this is particularly common in males. Ant queens are an important exception. The case of ant queen size dimorphisms has not been studied in sufficient detail, and thus this thesis aimed at elucidating causes and consequences of the different size of small (microgynous) and large (macrogynous)ant queens using the North American ant species Leptothorax rugatulus as a model system. Employing neutral genetic markers, no evidence for a taxonomically relevant separation of the gene pools of macrogynes and microgynes was found. Queens in polygynous colonies were highly related to each other, supporting the hypothesis that colonies with more than one queen commonly arise by secondary polygyny, i.e. by the adoption of daughter queens into their natal colonies. These results and conclusions are also true for the newly discovered queen size polymorphism in Leptothorax cf. andrei. Several lines of evidence favor the view that macrogynes predominantly found their colonies independently, while microgynes are specialized for dependent colony founding by readoption. Under natural conditions, mother and daughter size are highly correlated and this is also true for laboratory colonies. However, the size of developing queens is influenced by queens present in the colony. Comparing populations across the distribution range, it turns out that queen morphology (head width and ovariole number) is more differentiated among populations than worker morphology (coloration, multivariate size and shape), colony characteristics (queen and worker number per colony) or neutral genetic variation. Northern and southern populations differed consistently which indicates the possibility of two different species. The queen size dimorphism in L. rugatulus did neither influence the sex ratio produced by a colony, nor its ratio of workers to gynes. However, the sex ratio covaried strongly across populations with the average number of queens per colony in accordance with sex ratio theory. At the colony level, sex ratio could not be explained by current theory and a hypothesis at the colony-level was suggested. Furthermore, queen body size has no significant influence on the amount of reproductive skew among queens. Generally, the skew in L. rugatulus is low, and supports incomplete control models, rather than the classic skew models. In eight of fourteen mixed or microgynous colonies, the relative contributions of individual queens to workers, gynes and males were significantly different. This was mainly due to the fact that relative body size was negatively correlated with the ratio of gynes to workers produced. This supports the kin conflict over caste determination hypothesis which views microgyny as a selfish reproductive tactic.}, subject = {Ameisen}, language = {en} } @phdthesis{Ng2001, author = {Ng, Eva Yee Wah}, title = {How did Listeria monocytogenes become pathogenic?}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-1752}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2001}, abstract = {Listeriae are Gram positive, facultative, saprophytic bacteria capable of causing opportunistic infections in humans and animals. This thesis presents three separate lines of inquiries that can lead to the eventual convergence of a global view of Listeria as pathogen in the light of evolution, genomics, and function. First, we undertook to resolve the phylogeny of the genus Listeria with the goal of ascertaining insights into the evolution of pathogenic capability of its members. The phylogeny of Listeriae had not yet been clearly resolved due to a scarcity of phylogenetically informative characters within the 16S and 23S rRNA molecules. The genus Listeria contains six species: L. monocytogenes, L. ivanovii, L. innocua, L. seeligeri, L. welshimeri, and L. grayi; of these, L. monocytogenes and L. ivanovii are pathogenic. Pathogenicity is enabled by a 10-15Kb virulence gene cluster found in L. seeligeri, L. monocytogenes and L. ivanovii. The genetic contents of the virulence gene cluster loci, as well as some virulence-associated internalin loci were compared among the six species. Phylogenetic analysis based on a data set of nucleic acid sequences from prs, ldh, vclA, vclB, iap, 16S and 23S rRNA genes identified L. grayi as the ancestral branch of the genus. This is consistent with previous 16S and 23S rRNA findings. The remainder 5 species formed two groupings. One lineage represents L. monocytogenes and L. innocua, while the other contains L. welshimeri, L. ivanovii and L. seeligeri, with L. welshimeri forming the deepest branch within this group. Deletion breakpoints of the virulence gene cluster within L. innocua and L. welshimeri support the proposed tree. This implies that the virulence gene cluster was present in the common ancestor of L. monocytogenes, L. innocua, L. ivanovii, L. seeligeri and L. welshimeri; and that pathogenic capability has been lost in two separate events represented by L. innocua and L. welshimeri. Second, we attempted to reconstitute L. innocua of its deleted virulence gene cluster, in its original chromosomal location, from the L. monocytogenes 12 Kb virulence gene cluster. This turned out particularly difficult because of the limits of genetic tools presently available for the organism. The reconstitution was partially successful. The methods and approaches are presented, and all the components necessary to complete the constructs are at hand for both L. innocua and the parallel, positive control of L. monocytogenes mutant deleted of its virulence gene cluster. Third, the sequencing of the entire genome of L. monocytogenes EGDe was undertaken as part of an EU Consortium. Our lab was responsible for 10 per cent of the labor intensive gap-closure and annotation efforts, which I helped coordinate. General information and comparisons with sister species L. innocua and a close Gram positive relative Bacillus subtilis are presented in context. The areas I personally investigated, namely, sigma factors and stationary phase functions, are also presented. L. monocytogenes and L. innocua both possess surprisingly few sigma factors: SigA, SigB, SigH, SigL, and an extra-cytoplasmic function type sigma factor (SigECF). The stationary phase genes of L. monocytogenes is compared to the well-studied, complex, stationary phase networks of B. subtilis. This showed that while genetic competence functions may be operative in unknown circumstances, non-sporulating Listeria opted for very different approaches of regulation from B. subtilis. There is virtually no overlap of known, stationary phase genes between Listeria and Gram negative model organism E. coli.}, subject = {Listeria monocytogenes}, language = {en} } @phdthesis{Dornhaus2002, author = {Dornhaus, Anna}, title = {The role of communication in the foraging process of social bees}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-3468}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2002}, abstract = {In the various groups of social bees, different systems of communication about food sources occur. These communication systems are different solutions to a common problem of social insects: efficiently allocating the necessary number of workers first to the task of foraging and second to the most profitable food sources. The solution chosen by each species depends on the particular ecological circumstances as well as the evolutionary history of that species. For example, the outstanding difference between the bumble bee and the honey bee system is that honey bees can communicate the location of profitable food sources to nestmates, which bumble bees cannot. To identify possible selection pressures that could explain this difference, I have quantified the benefits of communicating location in honey bees. I show that these strongly depend on the habitat, and that communicating location might not benefit bees in temperate habitats. This could be due to the differing spatial distributions of resources in different habitats, in particular between temperate and tropical regions. These distributions may be the reason why the mostly temperate-living bumble bees have never evolved a communication system that allows them to transfer information on location of food sources, whereas most tropical social bees (all honey bees and many stingless bees) are able to recruit nestmates to specific points in their foraging range. Nevertheless, I show that in bumble bees the allocation of workers to foraging is also regulated by communication. Successful foragers distribute in the nest a pheromone which alerts other bees to the presence of food. This pheromone stems from a tergite gland, the function of which had not been identified previously. Usage of a pheromone in the nest to alert other individuals to forage has not been described in other social insects, and might constitute a new mode of communicating about food sources. The signal might be modulated depending on the quality of the food source. Bees in the nest sample the nectar that has been brought into the nest. Their decision whether to go out and forage depends not only on the pheromone signal, but also on the quality of the nectar they have sampled. In this way, foraging activity of a bumble bee colony is adjusted to foraging conditions, which means most bees are allocated to foraging only if high-quality food sources are available. In addition, foraging activity is adjusted to the amount of food already stored. In a colony with full honeypots, no new bees are allocated to foraging. These results help us understand how the allocation of workers to the task of food collection is regulated according to external and internal nest conditions in bumble bees.}, subject = {Hummel}, language = {en} } @article{PoethkeHovestadtMitesser2003, author = {Poethke, Hans-Joachim and Hovestadt, Thomas and Mitesser, Oliver}, title = {Local extinction and the evolution of dispersal rates: Causes and correlations}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-47718}, year = {2003}, abstract = {We present the results of individual-based simulation experiments on the evolution of dispersal rates of organisms living in metapopulations. We find conflicting results regarding the relationship between local extinction rate and evolutionarily stable (ES) dispersal rate depending on which principal mechanism causes extinction: if extinction is caused by environmental catastrophes eradicating local populations, we observe a positive correlation between extinction and ES dispersal rate; if extinction is a consequence of stochastic local dynamics and environmental fluctuations, the correlation becomes ambiguous; and in cases where extinction is caused by dispersal mortality, a negative correlation between local extinction rate and ES dispersal rate emerges. We conclude that extinction rate, which both affects and is affected by dispersal rates, is not an ideal predictor for optimal dispersal rates.}, subject = {Ausbreitung}, language = {en} } @phdthesis{Herzner2004, author = {Herzner, Gudrun}, title = {Evolution of the pheromone communication system in the European Beewolf Philanthus triangulum F. (Hymenoptera: Crabronidae)}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-11651}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2004}, abstract = {Darwin's theory of sexual selection explains the evolution of flamboyant male traits through female choice. It does not, however, address the question why males typically court and females choose. This asymmetry is now thought to be the result of the dichotomy in reproductive expenditures: Females invest primarily in parental care and males invest predominantly in mate attraction or competition. Based on this view, several hypotheses for the origin and maintenance of female preferences have been proposed. They include the classical sexual selection models, i.e. female choice for direct and indirect benefits as well as the more recent concepts of female choice for genetic compatibility and receiver bias models. The complementary choice scenario assumes that females choose mates with regard to genetic compatibility. The receiver bias concept views male traits and female preferences within the framework of communication theory and encompasses various more or less distinct models, two of which are sensory exploitation and sensory trap. Both models postulate that male signals evolved in response to pre-existing perceptual biases of females. The sensory trap hypothesis additionally emphasizes that pre-existing female preferences for certain cues evolved in non-sexual contexts, like e.g. foraging. Males that mimic these cues and elicit a favourable out-of-context response by females may increase their reproductive success. This thesis examines the evolution of the pheromone communication in the European Beewolf Philanthus triangulum. Beewolf females are specialized hunters of honeybees and provision their progeny with paralyzed prey. Male beewolves establish and scent mark territories with a pheromone from a head gland to court females. The concordant occurrence of the otherwise rare alcohol (Z)-11-eicosen-1-ol in the male pheromone and in the alarm pheromone of honeybees, the exclusive prey of the females, suggests a sensory trap process as an explanation for the evolution of the male pheromone in P. triangulum. According to this hypothesis, we tested three predictions: First, foraging honeybees should emit eicosenol. Via chemical analysis we could show that honeybee workers in fact smell of eicosenol during foraging. The occurrence of eicosenol on the cuticle and in the headspace of honeybees is a new finding. Second, beewolf females should use eicosenol as a cue for prey detection or identification. Using behavioural assays, we demonstrated that prey recognition in beewolf females is accomplished by olfactory cues and that eicosenol is an essential cue in this process. The sensory sensitivity of beewolf females to eicosenol must be extremely high, since they perceive the trace amounts present in the head space of honeybees. This sensitivity may be due to specialized olfactory receptors on the antennae of beewolf females. An inventory of the flagellar sensilla of both sexes showed that females carry one type of sensillum that is missing in males, the large sensillum basiconicum. This chemo-sensitive sensillum most likely plays a role in prey recognition. The third prediction is that beewolf males incorporate bee-like substances, including eicosenol, into their pheromone, and possibly catch females in a sensory trap. A reanalysis of the male pheromone revealed, among others, eicosenol and several alkanes and alkenes as pheromonal compounds. Our own analyses of the chemical profiles of honeybee workers and beewolf pheromone disclosed a surprisingly strong resemblance between the two. Eight of the eleven substances of the male pheromone are also present on the cuticle and in the headspace of honeybees. Notwithstanding this similarity, the male pheromone does not function as a sensory trap for females. Nevertheless, the extensive congruence between the odour bouquets of the females' prey and the male pheromone strongly suggests that the male signal evolved to exploit a pre-existing female sensory bias towards bee odour, and, thus represents a case of sensory exploitation. In addition to the above described scenario concerning mostly the 'design' of the male pheromone, we addressed possible indirect benefits female beewolves may gain by basing their mating decisions on signal 'content'. We show that the pheromone of male beewolves varies between families and may, thus, contain information about the degree of relatedness between the female and a potential mate. Females could use this information to choose genetically complementary males to avoid inbreeding and the production of infertile diploid sons. Collectively, our results provide strong evidence for a receiver bias process in the evolution of the male pheromone of P. triangulum. They further indicate that the pheromone composition may subsequently have been influenced by other natural or sexual selection pressures, like e.g. complementary female choice.}, subject = {Philantus}, language = {en} } @phdthesis{Kraft2004, author = {Kraft, Christian}, title = {Die Rolle von Mutation und Rekombination in der Mikroevolution von Helicobacter pylori}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-9757}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2004}, abstract = {Helicobacter pylori ist ein pathogenes Bakterium, das verantwortlich gemacht wird f{\"u}r verschiedene Erkrankungen des Magens und Duodenums, wie beispielsweise chronische Gastritis, peptische Ulzera und maligne Lymphome. Das Bakterium zeichnet sich durch eine hohe Rekombinationsrate aus und besitzt ein hohes Maß an genetischer Allelvielfalt. Im ersten Teil dieser Arbeit wurde die Rekombinationsrate und die L{\"a}nge der rekombinerten DNA-Importe anhand von sequentiellen Isolaten, die zu definierten Zeitpunkten aus dem selben Patienten isoliert wurden, untersucht. Es wurden zehn Gene, darunter sieben 'housekeeping' Gene und drei virulenzassoziierte Gene, amplifiziert und sequenziert. Die Ergebnisse zeigten eine bis dahin noch nicht f{\"u}r Bakterien beschriebene Fragmentl{\"a}nge der DNA-Importe von durchschnittlich lediglich 417 Basenpaaren. Die Rekombinationsrate war außergew{\"o}hnlich hoch. DNA-Microarray-Analysen konnten zeigen, dass es trotz dieser hohen Rekombinationsrate nur wenige Ver{\"a}nderungen in der genomischen Genausstattung gab. Jedoch hing das Auftreten von Rekombinationsereignissen direkt mit Ver{\"a}nderungen der Genausstattung zusammen. Im zweiten Teil der Arbeit wurde ein neues in vitro-Transformationsmodell entwickelt, das die in vivo ermittelten Resultate nachvollziehen sollte. Das Modell konnte sowohl die in vivo gefundene Rekombinationsrate als auch den Import von kurzen DNA-Fragmenten best{\"a}tigen, die zu einem Allelmosaik zwischen DNA-Rezipient und Donor f{\"u}hrten. Auff{\"a}llig war eine stark verminderte Transformierbarkeit mit Donor-DNA aus asiatischen H. pylori-St{\"a}mmen. Um eine m{\"o}gliche Beteiligung des Nukleotid-Excisions-Reparatur (NER) Mechanismus an der Rekombination zu ermitteln, wurden zwei Gene des Mechanismus ausgeschaltet. Die Ergebnisse der NER--Mutanten (uvrA-, uvrD-) zeigten eine starke Verminderung der Transformierbarkeit. Diese Verminderung hatte jedoch keinen Einfluss auf die L{\"a}nge der rekombinierten DNA-Importe. Das Ausschalten des uvrA-Gens f{\"u}hrte zudem zu einer erh{\"o}hten Sensibilit{\"a}t gegen{\"u}ber UV-Licht. Der NER-Mechanismus ist bei H. pylori in einer noch nicht aufgekl{\"a}rten Weise an der Rekombination beteiligt. In einem Rhesusaffen-Tiermodell wurde die initiale Besiedlung mit H. pylori untersucht. Die Tiere stellen einen nat{\"u}rlichen Wirt dar und zeigen {\"a}hnliche Krankheitssymptome wie menschliche Patienten. Die Rhesusaffen wurden experimentell mit zwei klinischen H. pylori-Isolaten infiziert. Die Reisolation zu bestimmten Zeitpunkten zeigte, dass sich nur einer der beiden St{\"a}mme im Affenmagen etablieren konnte und der zweite Stamm verdr{\"a}ngt worden war. In einem zweiten Versuchsansatz wurden die persistent infizierten Affen mit vier weiteren H. pylori-St{\"a}mmen infiziert, um eine transiente Koinfektion zu simulieren. Diese St{\"a}mme verdr{\"a}ngten jedoch den bereits etablierten Stamm, und es konnte keine in vivo-Rekombination festgestellt werden. Dennoch ist dieses Modell das Erste, in dem eine persistierende experimentelle H. pylori-Infektion in Rhesusaffen {\"u}ber einen Zeitraum von mehr als vier Jahren nachgewiesen werden konnte. Die Ergebnisse liefern wichtige Hinweise auf den beim Menschen meist unentdeckten Anfang der H. pylori-Infektion. Die Untersuchungen an weiteren Spezies des Genus Helicobacter zeigten, dass die beschriebene Spezies Heelicobacter nemestrinae keine eigene Spezies darstellt, sondern der Spezies H. pylori zugeordnet werden konnte. Den damit n{\"a}chsten 'Verwandten' stellt die Spezies H. acinonychis dar, deren St{\"a}mme sich untereinander wesentlich weniger stark unterscheiden als H. pylori-St{\"a}mme. Die Ergebnisse dieser Arbeit liefern wichtige Daten zum Verst{\"a}ndnis der Evolution und Mikroevolution innerhalb eines Wirtes von H. pylori, die zu besseren Strategien in der Bek{\"a}mpfung dieses pathogenen Bakteriums f{\"u}hren k{\"o}nnen.}, subject = {Helicobacter pylori}, language = {de} } @article{HuebnerWissemann2004, author = {H{\"u}bner, Stefanie and Wissemann, Volker}, title = {Morphometrische Analysen zur Variabilit{\"a}t von Prunus spinosa L. - Populationen(Prunoideae, Rosaceae) im mittleren Saaletal, Th{\"u}ringen}, doi = {10.3264/FG.2004.1215}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-35207}, year = {2004}, abstract = {Prunus spinosa L. (Rosaceae) is one of the most widespread members of the genus Prunus in middle europe. Its morphological plasticity resulted in a number of described taxa at subspecific level. Since the early neolithic times, drupes of the plum family are recorded and exhibit already a remarkable diversity in size and form. Here we present a short historical account to the use of P. spinosa and an overview of the different taxonomic treatments. We examined distribution patterns in general and in particular in the central valley of the river Saale (Thuringia) with respect to ecological, edaphic and climatic factors. We assessed within 16 populations the variability of 22 metric and 10 qualitative morphological characters at 7 different locations. Population sites included forest-, way- and fieldsides, as well as lightish pine forests. Pollen fertility did not increase during the flowering period, all flowers were directly fully fertile from the beginning. In contrast, glucose content varied significantly depending of the status of fertilization. Epicuticular wax structure was without variation amongst the populations. P. spinosa leaves are covered with a smooth layer of slightly striated wax. Morphological characters were scored on 270 branches and 506 fruits. Most of the characters showed enormous variability among and within populations such as metrics of leaves, thorns and character states of flower morphology. The lowest variability among populations and therefore not dependend of modificatory factors was found in fruit characters. Since kernel morphology seems to be genetically rather than modificatory controlled, we applied the 3 taxonomical concepts of Werneck, K{\"u}hn and Scholz u. Scholz to identify evolutionary units at subspecific levels. However, population variability was still so high, that from our study here we can not support an infraspecific classification of Prunus spinosa L.}, subject = {Prunus}, language = {de} } @phdthesis{Pils2005, author = {Pils, Birgit}, title = {Insights into the evolution of protein domains give rise to improvements of function prediction}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-16805}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2005}, abstract = {The growing number of uncharacterised sequences in public databases has turned the prediction of protein function into a challenging research field. Traditional annotation methods are often error-prone due to the small subset of proteins with experimentally verified function. Goal of this thesis was to analyse the function and evolution of protein domains in order to understand molecular processes in the cell. The focus was on signalling domains of little understood function, as well as on functional sites of protein domains in general. Glucosaminidases (GlcNAcases) represent key enzymes in signal transduction pathways. Together with glucosamine transferases, they serve as molecular switches, similar to kinases and phosphatases. Little was known about the molecular function and structure of the GlcNAcases. In this thesis, the GlcNAcases were identified as remote homologues of N-acetyltransferases. By comparing the homologous sequences, I was able to predict functional sites of the GlcNAcase family and to identify the GlcNAcases as the first family member of the acetyltransferase superfamily with a distinct catalytic mechanism, which is not involved in the transfer of acetyl groups. In a similar approach, the sensor domain of a plant hormone receptor was studied. I was able to predict putative ligand-binding sites by comparing evolutionary constraints in functionally diverged subfamilies. Most of the putative ligand-binding sites have been experimentally confirmed in the meantime. Due to the importance of enzymes involved in cellular signalling, it seems impossible to find substitutions of catalytic amino acids that turn them catalytically inactive. Nevertheless, by scanning catalytic positions of the protein tyrosine phosphatase families, I found many inactive domains among single domain and tandem domain phosphatases in metazoan proteomes. In addition, I found that inactive phosphatases are conserved throughout evolution, which led to the question about the function of these catalytically inactive phosphatase domains. An analysis of evolutionary site rates of amino acid substitutions revealed a cluster of conserved residues in the apparently redundant domain of tandem phosphatases. This putative regulatory center might be responsible for the experimentally verified dimerization of the active and inactive domain in order to control the catalytic activity of the active phosphatase domain. Moreover, I detected a subgroup of inactive phosphatases, which presumably functions in substrate recognition, based on different evolutionary site rates within the phosphatase family. The characterization of these new regulatory modules in the phosphatase family raised the question whether inactivation of enzymes is a more general evolutionary mechanism to enlarge signalling pathways and whether inactive domains are also found in other enzyme families. A large-scale analysis of substitutions at catalytic positions of enzymatic domains was performed in this work. I identified many domains with inactivating substitutions in various enzyme families. Signalling domains harbour a particular high occurrence of catalytically inactive domains indicating that these domains have evolved to modulate existing regulatory pathways. Furthermore, it was shown that inactivation of enzymes by single substitutions happened multiple times independently in evolution. The surprising variability of amino acids at catalytic positions was decisive for a subsequent analysis of the diversity of functional sites in general. Using functional residues extracted from structural complexes I could show that functional sites of protein domains do not only vary in their type of amino acid but also in their structural location within the domain. In the process of evolution, protein domains have arisen from duplication events and subsequently adapted to new binding partners and developed new functions, which is reflected in the high variability of functional sites. However, great differences exist between domain families. The analysis demonstrated that functional sites of nuclear domains are more conserved than functional sites of extracellular domains. Furthermore, the type of ligand influences the degree of conservation, for example ion binding sites are more conserved than peptide binding sites. The work presented in this thesis has led to the detection of functional sites in various protein domains involved in signalling pathways and it has resulted in insights into the molecular function of those domains. In addition, properties of functional sites of protein domains were revealed. This knowledge can be used in the future to improve the prediction of protein function and to identify functional sites of proteins.}, subject = {Dom{\"a}ne }, language = {en} } @phdthesis{Mitesser2006, author = {Mitesser, Oliver}, title = {The evolution of insect life history strategies in a social context}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-22576}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2006}, abstract = {This thesis extends the classical theoretical work of Macevicz and Oster (1976, expanded by Oster and Wilson, 1978) on adaptive life history strategies in social insects. It focuses on the evolution of dynamic behavioural patterns (reproduction and activity) as a consequence of optimal allocation of energy and time resources. Mathematical modelling is based on detailed empirical observations in the model species Lasioglossum malachurum (Halictidae; Hymenoptera). The main topics are field observations, optimisation models for eusocial life histories, temporal variation in life history decisions, and annual colony cycles of eusocial insects.}, subject = {Schmalbienen}, language = {en} } @article{GrosHovestadtPoethke2006, author = {Gros, Andreas and Hovestadt, Thomas and Poethke, Hans Joachim}, title = {Evolution of local adaptions in dispersal strategies}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-45406}, year = {2006}, abstract = {The optimal probability and distance of dispersal largely depend on the risk to end up in unsuitable habitat. This risk is highest close to the habitat's edge and consequently, optimal dispersal probability and distance should decline towards the habitat's border. This selection should lead to the emergence of spatial gradients in dispersal strategies. However, gene flow caused by dispersal itself is counteracting local adaptation. Using an individual based model we investigate the evolution of local adaptations of dispersal probability and distance within a single, circular, habitat patch. We compare evolved dispersal probabilities and distances for six different dispersal kernels (two negative exponential kernels, two skewed kernels, nearest neighbour dispersal and global dispersal) in patches of different size. For all kernels a positive correlation between patch size and dispersal probability emerges. However, a minimum patch size is necessary to allow for local adaptation of dispersal strategies within patches. Beyond this minimum patch area the difference in mean dispersal distance between center and edge increases linearly with patch radius, but the intensity of local adaptation depends on the dispersal kernel. Except for global and nearest neighbour dispersal, the evolved spatial pattern are qualitatively similar for both, mean dispersal probability and distance. We conclude, that inspite of the gene-flow originating from dispersal local adaptation of dispersal strategies is possible if a habitat is of sufficient size. This presumably holds for any realistic type of dispersal kernel.}, subject = {Ausbreitung}, language = {en} } @phdthesis{Baier2007, author = {Baier, Andrea}, title = {Architektur meiotischer Chromosomen : Eigenschaften und Evolution des Synaptonemalkomplexproteins SYCP3}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-25995}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2007}, abstract = {Die Meiose ist eine besondere Art der Zellteilung, die w{\"a}hrend der Keimzellreifung stattfindet. Sie umfasst zwei aufeinander folgende Zellteilungen mit nur einer DNA-Repli-kationsrunde, wodurch aus einer diploiden Ausgangszelle vier haploide Gameten entstehen. In der ersten meiotischen Teilung werden die homologen Chromosomen miteinander rekombiniert und voneinander getrennt, in der Meiose II findet die Trennung der Schwesterchromatiden statt. F{\"u}r den korrekten Ablauf dieser Prozesse musste sich eine spezielle molekulare Architektur des meiotischen Chromosoms entwickeln welche die Synapse der homologen Chromosomen durch den Synaptonemalkomplex (SC) beinhaltet. SCs sind evolution{\"a}r hochkonservierte, meiosespezifische Proteinkomplexe, die eine zentrale Bedeutung f{\"u}r Synapse, Rekombination und Segregation der homologen Chromosomen haben. Ein SC besteht aus zwei lateralen Elementen (LEs), die den Achsen der homologen Chromosomen aufgelagert sind, einer zentralen Region (CR) und einem zentralen Element (CE). Eine Hauptstrukturkomponente der LEs in Vertebraten ist das Synaptonemalkomplexprotein, SYCP3. Um die molekulare Architektur des SC besser zu verstehen und die Bedeutung von SYCP3 f{\"u}r die Zusammenlagerung der LE aufzudecken, wurden die Polymerisationseigenschaften von SYCP3, exprimiert in somatischen Zellen, erforscht. In diesem experimentellen Ansatz polymerisierte SYCP3 autonom zu stabilen, h{\"o}her geordneten, filament{\"o}sen Strukturen. Die „Coiled-Coil"-Dom{\"a}ne und die flankierenden, evolution{\"a}r konservierten Motive sind dabei notwenig, und nach Deletion des weniger konservierten N-terminalen Bereichs auch ausreichend f{\"u}r die Bildung der h{\"o}her geordneten Strukturen. Der N-Terminus hingegen spielt eine Rolle in der Stabilit{\"a}t der Polym{\"a}rstrukturen, welche durch Phosphorylierung zweier Serinreste im N-terminalen Bereich beeinflusst werden k{\"o}nnte. Obwohl die Struktur des SC in der Evolution hochkonserviert ist, sind die Protein-komponenten auf Aminos{\"a}uresequenzebene sehr unterschiedlich und weisen wenn {\"u}berhaupt eine strukturelle Homologie in ihrer Dom{\"a}nenorganisation auf. Um den SC-Aufbau und dessen Funktion besser verstehen zu k{\"o}nnen, wurden die orthologen SC-Proteine zwischen taxonomisch entfernten Spezies Ratte und Medaka verglichen. Es konnte gezeigt werden, dass trotz der Unterschiede in den Aminos{\"a}uresequenzen die sich in den letzen 450 Millionen Jahren zwischen Fisch- und S{\"a}ugern-SYCP3 akkumuliert haben, die Eigenschaften der Proteine vergleichbar sind, und das sie unter experimentellen Bedingungen miteinander interagieren und zu h{\"o}her geordneten Strukturen kopolymerisieren k{\"o}nnen.}, subject = {Meiose}, language = {de} } @unpublished{Dandekar2007, author = {Dandekar, Thomas}, title = {Some general system properties of a living observer and the environment he explores}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-33537}, year = {2007}, abstract = {In a nice assay published in Nature in 1993 the physicist Richard God III started from a human observer and made a number of witty conclusions about our future prospects giving estimates for the existence of the Berlin Wall, the human race and all the rest of the universe. In the same spirit, we derive implications for "the meaning of life, the universe and all the rest" from few principles. Adams´ absurd answer "42" tells the lesson "garbage in / garbage out" - or suggests that the question is non calculable. We show that experience of "meaning" and to decide fundamental questions which can not be decided by formal systems imply central properties of life: Ever higher levels of internal representation of the world and an escalating tendency to become more complex. An observer, "collecting observations" and three measures for complexity are examined. A theory on living systems is derived focussing on their internal representation of information. Living systems are more complex than Kolmogorov complexity ("life is NOT simple") and overcome decision limits (G{\"o}del theorem) for formal systems as illustrated for cell cycle. Only a world with very fine tuned environments allows life. Such a world is itself rather complex and hence excessive large in its space of different states - a living observer has thus a high probability to reside in a complex and fine tuned universe.}, subject = {Komplex }, language = {en} } @article{MitesserWeisselStrohmetal.2007, author = {Mitesser, Oliver and Weissel, Norbert and Strohm, Erhard and Poethke, Hans-Joachim}, title = {Adaptive dynamic resource allocation in annual eusocial insects: Environmental variation will not necessarily promote graded control}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-45412}, year = {2007}, abstract = {Background: According to the classical model of Macevicz and Oster, annual eusocial insects should show a clear dichotomous "bang-bang" strategy of resource allocation; colony fitness is maximised when a period of pure colony growth (exclusive production of workers) is followed by a single reproductive period characterised by the exclusive production of sexuals. However, in several species graded investment strategies with a simultaneous production of workers and sexuals have been observed. Such deviations from the "bang-bang" strategy are usually interpreted as an adaptive (bet-hedging) response to environmental fluctuations such as variation in season length or food availability. To generate predictions about the optimal investment pattern of insect colonies in fluctuating environments, we slightly modified Macevicz and Oster's classical model of annual colony dynamics and used a dynamic programming approach nested into a recurrence procedure for the solution of the stochastic optimal control problem. Results: 1) The optimal switching time between pure colony growth and the exclusive production of sexuals decreases with increasing environmental variance. 2) Yet, for reasonable levels of environmental fluctuations no deviation from the typical bang-bang strategy is predicted. 3) Model calculations for the halictid bee Lasioglossum malachurum reveal that bet-hedging is not likely to be the reason for the graded allocation into sexuals versus workers observed in this species. 4) When environmental variance reaches a critical level our model predicts an abrupt change from dichotomous behaviour to graded allocation strategies, but the transition between colony growth and production of sexuals is not necessarily monotonic. Both, the critical level of environmental variance as well as the characteristic pattern of resource allocation strongly depend on the type of function used to describe environmental fluctuations. Conclusion: Up to now bet-hedging as an evolutionary response to variation in season length has been the main argument to explain field observations of graded resource allocation in annual eusocial insect species. However, our model shows that the effect of moderate fluctuations of environmental conditions does not select for deviation from the classical bang-bang strategy and that the evolution of graded allocation strategies can be triggered only by extreme fluctuations. Detailed quantitative observations on resource allocation in eusocial insects are needed to analyse the relevance of alternative explanations, e.g. logistic colony growth or reproductive conflict between queen and workers, for the evolution of graded allocation strategies.}, subject = {Insekten}, language = {en} } @phdthesis{Zdziarski2008, author = {Zdziarski, Jaroslaw Maciej}, title = {Bacterial Genome Plasticity and its Role for Adaptation and Evolution of Asymptomatic Bacteriuria (ABU) Escherichia coli Strains}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-32879}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2008}, abstract = {Asymptomatic bacteriuria (ABU) represents the long term bacterial colonization of the urinary tract, frequently caused by Escherichia coli (E. coli), without typical symptoms of a urinary tract infection (UTI). To investigate characteristics of ABU E. coli isolates in more detail, the geno- and phenotypes of eleven ABU isolates have been compared. Moreover, consecutive in vivo re-isolates of the model ABU strain 83972 were characterized with regard to transcriptomic, proteomic and genomic alterations upon long term in vivo persistence in the human bladder. Finally, the effect of the human host on bacterial adaptation/evolution was assessed by comparison of in vitro and in vivo-propagated strain 83972. ABU isolates represent a heterologous group of organisms. The comparative analysis of different ABU isolates elucidated the remarkable genetic and phenotypic flexibility of E. coli isolates. These isolates could be allocated to all four major E. coli phylogenetic lineages as well as to different clonal groups. Accordingly, they differed markedly in genome content, i.e., the genome size as well as the presence of typical UPEC virulence-associated genes. Multi locus sequence typing suggested that certain ABU strains evolved from UPEC variants that are able to cause symptomatic UTI by genome reduction. Consequently, the high E. coli genome plasticity does not allow a generalized view on geno- and phenotypes of individual isolates within a clone. Reductive evolution by point mutations, DNA rearrangements and deletions resulted in inactivation of genes coding for several UPEC virulence factors, thus supporting the idea that a reduced bacterial activation of host mucosal inflammation promotes the ABU lifestyle of these E. coli isolates. Gene regulation and genetic diversity are strategies which enable bacteria to live and survive under continuously changing environmental conditions. To study adaptational changes upon long term growth in the bladder, consecutive re-isolates of model ABU strain 83972 derived from a human colonisation study and from an in vitro long term cultivation experiment were analysed with regard to transcriptional changes and genome rearrangements. In this context, it could be demonstrated that E. coli, when exposed to different host backgrounds, is able to adapt its metabolic networks resulting in an individual bacterial colonisation strategy. Transcriptome and proteome analyses demonstrated distinct metabolic strategies of nutrients acquisition and energy production of tested in vivo re-isolates of strain 83972 that enabled them to colonise their host. Utilisation of D-serine, deoxy- and ribonucleosides, pentose and glucuronate interconversions were main up-regulated pathways providing in vivo re-isolates with extra energy for efficient growth in the urinary bladder. Moreover, this study explored bacterial response networks to host defence mechanisms: The class III alcohol dehydrogenase AdhC, already proven to be involved in nitric oxide detoxification in pathogens like Haemophilus influenzae, was shown for the first time to be employed in defending E. coli against the host response during asymptomatic bacteriuria. Consecutive in vivo and in vitro re-isolates of strain 83972 were also analysed regarding their genome structure. Several changes in the genome structure of consecutive re-isolates derived from the human colonisation study implied the importance of bacterial interactions with the host during bacterial microevolution. In contrast, the genome structure of re-isolates from the in vitro long term cultivation experiment, where strain 83972 has been propagated without host contact, was not affected. This suggests that exposure to the immune response promotes genome plasticity thus being a driving force for the development of the ABU lifestyle and evolution within the urinary tract.}, subject = {Escherichia coli}, language = {en} } @phdthesis{Pinkert2008, author = {Pinkert, Stefan}, title = {The human proteome is shaped by evolution and interactions}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-35566}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2008}, abstract = {Das menschliche Genom ist seit 2001 komplett sequenziert. Ein Großteil der Proteine wurde mittlerweile beschrieben und t{\"a}glich werden bioinformatische Vorhersagen praktisch best{\"a}tigt. Als weiteres Großprojekt wurde k{\"u}rzlich die Sequenzierung des Genoms von 1000 Menschen gestartet. Trotzdem ist immer noch wenig {\"u}ber die Evolution des gesamten menschlichen Proteoms bekannt. Proteindom{\"a}nen und ihre Kombinationen sind teilweise sehr detailliert erforscht, aber es wurden noch nicht alle Dom{\"a}nenarchitekturen des Menschen in ihrer Gesamtheit miteinander verglichen. Der verwendete große hochqualitative Datensatz von Protein-Protein-Interaktionen und Komplexen stammt aus dem Jahr 2006 und erm{\"o}glicht es erstmals das menschliche Proteom mit einer vorher nicht m{\"o}glichen Genauigkeit analysieren zu k{\"o}nnen. Hochentwickelte Cluster Algorithmen und die Verf{\"u}gbarkeit von großer Rechenkapazit{\"a}t bef{\"a}higen uns neue Information {\"u}ber Proteinnetzwerke ohne weitere Laborarbeit zu gewinnen. Die vorliegende Arbeit analysiert das menschliche Proteom auf drei verschiedenen Ebenen. Zuerst wurde der Ursprung von Proteinen basierend auf ihrer Dom{\"a}nenarchitektur analysiert, danach wurden Protein-Protein-Interaktionen untersucht und schließlich erfolgte Einteilung der Proteine nach ihren vorhandenen und fehlenden Interaktionen. Die meisten bekannten Proteine enthalten mindestens eine Dom{\"a}ne und die Proteinfunktion ergibt sich aus der Summe der Funktionen der einzelnen enthaltenen Dom{\"a}nen. Proteine, die auf der gleichen Dom{\"a}nenarchitektur basieren, das heißt die die gleichen Dom{\"a}nen in derselben Reihenfolge besitzen, sind homolog und daher aus einem gemeinsamen urspr{\"u}nglichen Protein entstanden. Die Dom{\"a}nenarchitekturen der urspr{\"u}nglichen Proteine wurden f{\"u}r 750000 Proteine aus 1313 Spezies bestimmt. Die Gruppierung von Spezies und ihrer Proteine ergibt sich aus taxonomischen Daten von NCBI-Taxonomy, welche mit zus{\"a}tzlichen Informationen basierend auf molekularen Markern erg{\"a}nzt wurden. Der resultierende Datensatz, bestehend aus 5817 Dom{\"a}nen und 32868 Dom{\"a}nenarchitekturen, war die Grundlage f{\"u}r die Bestimmung des Ursprungs der Proteine aufgrund ihrer Dom{\"a}nenarchitekturen. Es wurde festgestellt, dass nur ein kleiner Teil der neu evolvierten Dom{\"a}nenarchitekturen eines Taxons gleichzeitig auch im selben Taxon neu entstandene Proteindom{\"a}nen enth{\"a}lt. Ein weiteres Ergebnis war, dass Dom{\"a}nenarchitekturen im Verlauf der Evolution l{\"a}nger und komplexer werden, und dass so verschiedene Organismen wie der Fadenwurm, die Fruchtfliege und der Mensch die gleiche Menge an unterschiedlichen Proteinen haben, aber deutliche Unterschiede in der Anzahl ihrer Dom{\"a}nenarchitekturen aufweisen. Der zweite Teil besch{\"a}ftigt sich mit der Frage wie neu entstandene Proteine Bindungen mit dem schon bestehenden Proteinnetzwerk eingehen. In fr{\"u}heren Arbeiten wurde gezeigt, dass das Protein-Interaktions-Netzwerk ein skalenfreies Netz ist. Skalenfreie Netze, wie zum Beispiel das Internet, bestehen aus wenigen Knoten mit vielen Interaktionen, genannt Hubs, und andererseits aus vielen Knoten mit wenigen Interaktionen. Man vermutet, dass zwei Mechanismen zur Entstehung solcher Netzwerke f{\"u}hren. Erstens m{\"u}ssen neue Proteine um auch Teil des Proteinnetzwerkes zu werden mit Proteinen interagieren, die bereits Teil des Netzwerkes sind. Zweitens interagieren die neuen Proteine, gem{\"a}ß der Theorie der bevorzugten Bindung, mit h{\"o}herer Wahrscheinlichkeit mit solchen Proteinen im Netzwerk, die schon an zahlreichen weiteren Protein-Interaktionen beteiligt sind. Die Human Protein Reference Database stellt ein auf Informationen aus in-vivo Experimenten beruhendes Proteinnetzwerk f{\"u}r menschliche Proteine zur Verf{\"u}gung. Basierend auf den in Kapitel I gewonnenen Informationen wurden die Proteine mit dem Ursprungstaxon ihrer Dom{\"a}nenarchitekturen versehen. Dadurch wurde gezeigt, dass ein Protein h{\"a}ufiger mit Proteinen, die im selben Taxon entstanden sind, interagiert, als mit Proteinen, die in anderen Taxa neu aufgetreten sind. Es stellte sich heraus, dass diese Interaktionsraten f{\"u}r alle Taxa deutlich h{\"o}her waren, als durch das Zufallsmodel vorhergesagt wurden. Alle Taxa enthalten den gleichen Anteil an Proteinen mit vielen Interaktionen. Diese zwei Ergebnisse sprechen dagegen, dass die bevorzugte Bindung der alleinige Mechanismus ist, der zum heutigen Aufbau des menschlichen Proteininteraktion-Netzwerks beigetragen hat. Im dritten Teil wurden Proteine basierend auf dem Vorhandensein und der Abwesenheit von Interaktionen in Gruppen eingeteilt. Proteinnetzwerke k{\"o}nnen in kleine hoch vernetzte Teile zerlegt werden, die eine spezifische Funktion aus{\"u}ben. Diese Gruppen k{\"o}nnen mit hoher statistischer Signifikanz berechnet werden, haben meistens jedoch keine biologische Relevanz. Mit einem neuen Algorithmus, welcher zus{\"a}tzlich zu Interaktionen auch Nicht-Interaktionen ber{\"u}cksichtigt, wurde ein Datensatz bestehend aus 8,756 Proteinen und 32,331 Interaktionen neu unterteilt. Eine Einteilung in elf Gruppen zeigte hohe auf Gene Ontology basierte Werte und die Gruppen konnten signifikant einzelnen Zellteilen zugeordnet werden. Eine Gruppe besteht aus Proteinen, welche wenige Interaktionen miteinander aber viele Interaktionen zu zwei benachbarten Gruppen besitzen. Diese Gruppe enth{\"a}lt eine signifikant erh{\"o}hte Anzahl an Transportproteinen und die zwei benachbarten Gruppen haben eine erh{\"o}hte Anzahl an einerseits extrazellul{\"a}ren und andererseits im Zytoplasma und an der Membran lokalisierten Proteinen. Der Algorithmus hat damit unter Beweis gestellt das die Ergebnisse nicht bloß statistisch sondern auch biologisch relevant sind. Wenn wir auch noch weit vom Verst{\"a}ndnis des Ursprungs der Spezies entfernt sind, so hat diese Arbeit doch einen Beitrag zum besseren Verst{\"a}ndnis der Evolution auf dem Level der Proteine geleistet. Im Speziellen wurden neue Erkenntnisse {\"u}ber die Beziehung von Proteindom{\"a}nen und Dom{\"a}nenarchitekturen, sowie ihre Pr{\"a}ferenzen f{\"u}r Interaktionspartner im Interaktionsnetzwerk gewonnen.}, subject = {Evolution}, language = {en} } @phdthesis{Gros2008, author = {Gros, Andreas}, title = {Interactions in the evolution of dispersal distance and emigration probability}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-29226}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2008}, abstract = {Chapter 1 - Evolution of local adaptations in dispersal strategies The optimal probability and distance of dispersal largely depend on the risk to end up in unsuitable habitat. This risk is highest close to the habitat's edge and consequently, optimal dispersal probability and distance should decline towards the habitat's border. This selection should lead to the emergence of spatial gradients in dispersal strategies. However, gene flow caused by dispersal itself is counteracting local adaptation. Using an individual based model I investigate the evolution of local adaptations of dispersal probability and distance within a single, circular, habitat patch. I compare evolved dispersal probabilities and distances for six different dispersal kernels (two negative exponential kernels, two skewed kernels, nearest neighbour dispersal and global dispersal) in patches of different size. For all kernels a positive correlation between patch size and dispersal probability emerges. However, a minimum patch size is necessary to allow for local adaptation of dispersal strategies within patches. Beyond this minimum patch area the difference in mean dispersal distance between center and edge increases linearly with patch radius, but the intensity of local adaptation depends on the dispersal kernel. Except for global and nearest neighbour dispersal, the evolved spatial pattern are qualitatively similar for both, mean dispersal probability and distance. I conclude, that inspite of the gene-flow originating from dispersal local adaptation of dispersal strategies is possible if a habitat is of sufficient size. This presumably holds for any realistic type of dispersal kernel. Chapter 2 - How dispersal propensity and distance depend on the capability to assess population density We analyze the simultaneous evolution of emigration probability and dispersal distance for species with different abilities to assess habitat quality (population density) and which suffer from distance dependent dispersal costs. Using an individual-based model I simulate dispersal as a multistep (patch to patch) process in a world consisting of habitat patches surrounded by lethal matrix. Our simulations show that natal dispersal is strongly driven by kin-competition but that consecutive dispersal steps are mostly determined by the chance to immigrate into patches with lower population density. Consequently, individuals following an informed strategy where emigration probability depends on local population density disperse over larger distances than individuals performing density-independent emigration; this especially holds when variation in environmental conditions is spatially correlated. However, already moderate distance-dependent dispersal costs prevent the evolution of long-distance dispersal irrespectively of the chosen dispersal strategy. Chapter 3 - Evolution of sex-biased dispersal: the role of sex-specific dispersal costs, demographic stochasticity, and inbreeding Inbreeding avoidance and asymmetric competition over resources have both been identified as factors favouring the evolution of sex- biased dispersal. It has also been recognized that sex-specific costs of dispersal would promote selection for sexspecific dispersal, but there is little quantitative information on this aspect. In this paper I explore (i) the quantitative relationship between cost-asymmetry and a bias in dispersal, (ii) the influence of demographic stochasticity on this effect, and (iii) how inbreeding and cost-asymmetry interact in their effect on sex-specific dispersal. I adjust an existing analytical model to account for sex-specific costs of dispersal. Based on numerical calculations I predict a severe bias in dispersal already for small differences in dispersal costs. I corroborate these predictions in individualbased simulations, but show that demographic stochasticity generally leads to more balanced dispersal. In combination with inbreeding, cost asymmetries will usually determine which of the two sexes becomes the more dispersive. Chapter 4 - Evolution of sex-biased dispersal: the role of sex-specific dispersal costs, demographic stochasticity, and inbreeding Inbreeding depression, asymmetries in costs or benefits, and the mating system have been identified as potential factors underlying the evolution of sex-biased dispersal. We use individual-based simulations to explore how the mating system and demographic stochasticity influence the evolution of sex-specific dispersal in a metapopulation with females competing over breeding sites, and males over mating opportunities. Comparison of simulation results for random mating with those for a harem system (locally, a single male sires all offspring) reveal that even extreme variance in local male reproductive success (extreme male competition) does not induce a male bias in dispersal. The latter evolves if between-patch variance in reproductive success is larger for males than females. This can emerge due to demographic stochasticity if habitat patches are small. More generally, members of a group of individuals experiencing higher spatio-temporal variance in fitness expectations may evolve to disperse with greater probability than others.}, subject = {Theoretische {\"O}kologie}, language = {en} } @phdthesis{Winkel2009, author = {Winkel, Karoline}, title = {Synaptonemalkomplexprotein SYCP1: Bindungspartner, Polymerisationseigenschaften und evolution{\"a}re Aspekte}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-43955}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2009}, abstract = {Synaptonemal Komplexe (SC) sind evolution{\"a}r konservierte, meiosespezifische, protein{\"o}se Strukturen, die maßgeblich an Synapsis, Rekombination und Segregation der homologen Chromosomen beteiligt sind. Sie zeigen eine dreigliedrige strickleiter-artige Organisation, die sich aus i) zwei Lateralelementen (LE), an die das Chromatin der Homologen angelagert ist, ii) zahlreichen Transversalfilamenten (TF), welche die LE in einer reißverschlussartigen Weise miteinander verkn{\"u}pfen, und iii) einem zentralen Element (CE) zusammensetzt. Die Hauptproteinkomponenten der S{\"a}uger-SC sind das Transversalfilamentprotein SYCP1 und die Lateralelementproteine SYCP2 und SYCP3. Wie sich die SC-Struktur zusammenf{\"u}gt war bisher nur wenig verstanden; es war nicht bekannt wie die TF innerhalb der LE-Strukturen verankert sind und dabei die homologen Chromosomen verkn{\"u}pfen. Aufgrund dessen wurde die Interaktion zwischen den Proteinen SYCP1 und SYCP2 untersucht. Mit der Hilfe verschiedenster Interaktionssysteme konnte gezeigt werden, dass der C-Terminus von SYCP1 mit SYCP2 interagieren kann. Aufgrund der Bindungsf{\"a}higkeit zu beiden Proteinen, SYCP1 und SYCP3, kann angenommen werden, dass SYCP2 als Linker zwischen diesen Proteinen fungiert und somit m{\"o}glicherweise das fehlende Bindungsglied zwischen den Lateralelementen und Transversalfilamenten darstellt. Obwohl die SC-Struktur in der Evolution hochkonserviert ist, schien dies nicht f{\"u}r seine Protein-Untereinheiten zuzutreffen. Um die Struktur und Funktion des SC besser verstehen zu k{\"o}nnen, wurde ein Vergleich zwischen den orthologen SYCP1 Proteinen der evolution{\"a}r entfernten Spezies Ratte und Medaka erstellt. Abgesehen von den erheblichen Sequenzunterschieden die sich in 450 Millionen Jahren der Evolution angeh{\"a}uft haben, traten zwei bisher nicht identifizierte Sequenzmotive hervor, CM1 und CM2, die hochgradig konserviert sind. Anhand dieser Motive konnte in Datenbankanalysen erstmals ein Protein in Hydra vulgaris nachgewiesen werden, bei dem es sich um das orthologe Protein von SYCP1 handeln k{\"o}nnte. Im Vergleich mit dem SYCP1 der Ratte zeigten die Proteine aus Medaka und Hydra, neben den hoch konservierten CM1 und CM2, vergleichbare Dom{\"a}nenorganisationen und im heterologen System zudem sehr {\"a}hnliche Polymerisationseigenschaften. Diese Ergebnisse sprechen f{\"u}r eine evolution{\"a}re Konservierung von SYCP1.}, subject = {Meiose}, language = {de} } @misc{OPUS4-3424, title = {Blick - das Magazin der Julius-Maximilians-Universit{\"a}t W{\"u}rzburg. Ausgabe 1/2009. Schwerpunktthema: Am Anfang war Charles Darwin}, volume = {1/2009}, organization = {Julius-Maximilians-Universit{\"a}t W{\"u}rzburg}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-40693}, year = {2009}, abstract = {Inhalts{\"u}bersicht zum Schwerpunktthema: - Leidenschaftlicher Sammler und Beobachter: Charles Darwin und die Entwicklung der Evolutionstheorie - Die Tumorzelle hat den Vorteil: Wo der Krebsforscher Manfred Schartl auf Spuren der Evolution st{\"o}ßt - Brillentr{\"a}ger sind die Zukunft: Der Humangenetiker Holger H{\"o}hn ist {\"u}berzeugt davon, dass die Evolution trotz moderner Medizin noch immer ihr Spiel treibt mit Homo sapiens - Wenn der Putzer zubeißt: Eine Vorlesungsstunde im Biozentrum {\"u}ber Evolution - Die Evolution als Rechenaufgabe: Am Lehrstuhl f{\"u}r Bioinformatik arbeiten Biologen, Mathematiker und Informatiker daran, den Stammbaum des Lebens zu berechnen - Am Anfang war das Wort - oder doch der Urknall?: Religi{\"o}se Wahrheit und naturwissenschaftliche Erkenntnis schließen sich gegenseitig nicht aus - Bruch beim gr{\"u}nen Heinrich: Was die Evolutionstheorie bei den Literaten ausl{\"o}ste - Auf Spurensuche im Vatikan: Ein Kirchengeschichtler untersucht die Reaktion der katholischen Kirche auf Darwins Evolutionstheorie u. a.}, subject = {W{\"u}rzburg}, language = {de} } @phdthesis{Vershenya2010, author = {Vershenya, Stanislav}, title = {Quantitative and qualitative analyses of in-paralogs}, url = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-51358}, school = {Universit{\"a}t W{\"u}rzburg}, year = {2010}, abstract = {In our analysis I was interested in the gene duplications, with focus on in-paralogs. In-paralogs are gene duplicates which arose after species split. Here I analysed the in-paralogs quantitatively, as well as qualitatively. For quantitative analysis genomes of 21 species were taken. Most of them have vastly different lifestyles with maximum evolutionary distance between them 1100 million years. Species included mammals, fish, insects and worm, plus some other chordates. All the species were pairwised analysed by the Inparanoid software, and in-paralogs matrix were built representing number of in-paralogs in all vs. all manner. Based on the in-paralogs matrix I tried to reconstruct the evolutionary tree using in-paralog numbers as evolutionary distance. If all 21 species were used the resulting tree was very far from real one: a lot of species were misplaced. However if the number was reduced to 12, all of the species were placed correctly with only difference being wrong insect and fish clusters switched. Then to in-paralogs matrix the neighbour-net algorithm was applied. The resulting "net" tree showed the species with fast or slow duplications rates compared to the others. We could identify species with very high or very low duplications frequencies and it correlates with known occurrences of the whole genome duplications. As the next step I built the graphs for every single species showing the correlation between their in-paralogs number and evolutionary distance. As we have 21 species, graph for every species is built using 20 points. Coordinates of the points are set using the evolutionary distance to that particular species and in-paralogs number. In mammals with increasing the distance from speciation the in-paralogs number also increased, however not in linear fashion. In fish and insects the graph close to zero is just the same in mammals' case. However, after reaching the evolutionary distances more than 800 million years the number of inparalogs is beginning to decrease. We also made a simulation of gene duplications for all 21 species and all the splits according to the fossil and molecular clock data from literature. In our simulation duplication frequency was minimal closer to the past and maximum in the near-present time. Resulting curves had the same shape the experimental data ones. In case of fish and insect for simulation the duplication rate coefficient even had to be set negative in order to repeat experimental curve shape. To the duplication rate coefficient in our simulation contribute 2 criteria: gene duplications and gene losses. As gene duplication is stochastical process it should always be a constant. So the changing in the coefficient should be solely explained by the increasing gene loss of old genes. The processes are explained by the evolution model with high gene duplication and loss ratio. The drop in number of in-paralogs is probably due to the BLAST algorithm. It is observed in comparing highly divergent species and BLAST cannot find the orthologs so precisely anymore. In the second part of my work I concentrated more on the specific function of inparalogs. Because such analysis is time-consuming it could be done on the limited number species. Here I used three insects: Drosophila melanogaster (fruit y), Anopheles gambiae (mosquito) and Apis mellifera (honeybee). After Inparnoid analyses and I listed the cluster of orthologs. Functional analyses of all listed genes were done using GO annotations and also KEGG PATHWAY database. We found, that the gene duplication pattern is unique for each species and that this uniqueness is rejected through the differences in functional classes of duplicated genes. The preferences for some classes reject the evolutionary trends of the last 350 million years and allow assumptions on the role of those genes duplications in the lifestyle of species. Furthermore, the observed gene duplications allowed me to find connections between genomic changes and their phenotypic manifestations. For example I found duplications within carbohydrate metabolism rejecting feed pattern adaptation, within photo- and olfactory-receptors indicating sensing adaptation and within troponin indicating adaptations in the development. Despite these species specific differences, found high correlations between the independently duplicated genes between the species. This might hint for a "pool" of genes preferentially duplicated. Taken together, the observed duplication patterns reject the adaptational process and provide us another link to the field of genomic zoology.}, subject = {Duplikation}, language = {en} }