Sostdc1 Plays an Essential Role in Mammalian Tooth Patterning : Insight into the Rodent Dental Evolution

This thesis work focuses on the role of TGF-beta family antagonists during the development of mouse dentition. Tooth develops through an interaction between the dental epithelium and underlying neural crest derived mesenchyme. The reciprocal signaling between these tissues is mediated by soluble signaling molecules and the balance between activatory and inhibitory signals appears to be essential for the pattern formation. We showed the importance of Sostdc1 in the regulation of tooth shape and number. The absence of Sostdc1 altered the molar cusp patterning and led to supernumerary tooth formation both in the molar and incisor region. We showed that initially, Sostdc1 expression is in the mesenchyme, suggesting that dental mesenchyme may limit supernumerary tooth induction. We tested this in wild-type incisors by minimizing the amount of mesenchymal tissue surrounding the incisor tooth germs prior to culture in vitro. The cultured teeth phenocopied the extra incisor phenotype of the Sostdc1-deficient mice. Furthermore, we showed that minimizing the amount of dental mesenchyme in cultured Sostdc1-deficient incisors caused the formation of additional de novo incisors that resembled the successional incisor development resulting from activated Wnt signaling. Sostdc1 seemed to be able to inhibit both mesenchymal BMP4 and epithelial canonical Wnt signaling, which thus allows Sostdc1 to restrict the enamel knot size and regulate the tooth shape and number. Our work emphasizes the dual role for the tooth mesenchyme as a suppressor as well as an activator during tooth development. We found that the placode, forming the thick mouse incisor, is prone to disintegration during initiation of tooth development. The balance between two mesenchymal TGF-beta family signals, BMP4 and Activin is essential in this regulation. The inhibition of BMP4 or increase in Activin signaling led to the splitting of the large incisor placode into two smaller placodes resulting in thin incisors. These two signals appeared to have different effects on tooth epithelium and the analysis of the double null mutant mice lacking Sostdc1 and Follistatin indicated that these TGF-beta inhibitors regulate the mutual balance of BMP and Activin in vivo. In addition, this work provides an alternative explanation for the issue of incisor identity published in Science by Tucker et al. in 1998 and proposes that the molar like morphology that can be obtained by inhibiting BMP signaling is due to partial splitting of the incisor placodes and not due to change in tooth identity from the incisor to the molar. This thesis work presents possible molecular mechanisms that may have modified the mouse dental pattern during evolution leading to the typical rodent dentition of modern mouse. The rodent dentition is specialized for gnawing and consists of two large continuously growing incisors and toothless diastema region separating the molars and incisors. The ancestors of rodents had higher number of more slender incisors together with canines and premolars. Additionally, murine rodents, which include the mouse, have lost their ability for tooth replacement. This work has revealed that the inhibitory molecules appear to play a role in the tooth number suppression by delineating the spatial and temporal action of the inductive signals. The results suggest that Sostdc1 plays an essential role in several stages of tooth development through the regulation of both the BMP and Wnt pathway. The work shows a dormant sequential tooth forming potential present in wild type mouse incisor region and gives a new perspective on tooth suppression by dental mesenchyme. It reveals as well a novel mechanism to create a large mouse incisor through the regulation of mesenchymal balance between inductive and inhibitory signals.

Population structure and evolution in the ant Plagiolepis pygmaea and its two social parasites Plagiolepis xene and Plagiolepis grassei

Social behaviour affects dispersal of animals and is an important modifier of genetic population structures. The female sex is often philopatric, which maintains coancestry within the breeding groups and promotes cooperative behaviours. This enables also inclusive fitness returns from altruism and explains why some individuals sacrifice personal reproduction for the good of others in social insects such as ants. However, reduced dispersal and population substructuring at the level of colonies may also entail inbreeding, loss of genetic diversity, and vulnerability. In addition, the most vulnerable ants are species that are evolved to parasitize colonies of other ants, and which compromise between abilities to disperse and the efficiency to parasitize the host. On the other hand, certain social organisations of ant colonies may facilitate a species to disperse outside its natural range and become a pest. Altogether, knowledge on genetic structuring of ant populations, as well as the evolution of their life histories can contribute to conservation biology and population management. The aim of this thesis was to investigate population structures and phylogenetic evolution of the ant Plagiolepis pygmaea and its two obligatory, workerless social parasites (inquilines) P. xene and P. grassei with genetic markers and DNA sequence data. The results support the general assumption that populations of inquiline parasites are highly fragmented and genetically vulnerable. Comparison of the two parasites suggests that differences in their relative abundance may follow from their interaction with the host, i.e. how well the species is adapted to reproduce in the host colonies. The results also indicate that the most recent free living ancestor to these two parasite species is their common host. This is considered to provide evidence for the controversial issue of sympatric speciation. Further, given that the level of adaptations to parasitic life history depends on the evolutionary time since the free-living ancestor, the results establish a link between species rarity and its evolutionary age. The populations of the host species P. pygmaea displayed significantly reduced dispersal both among the females (queens) and males, and high levels of inbreeding which may enhance worker altruism. In addition, the queens were found to mate with multiple males. Given the high relatedness between the queens and their mates, this occurs probably for non-genetic reasons, e.g. without benefits associated in genetically more diverse offspring. The results hence caution that the contribution of non-genetic factors to the prevailing mating patterns and genetic population structures should not be underestimated.

Effects of turbidity on feeding and distribution of fish

In aquatic systems, the ability of both the predator and prey to detect each other may be impaired by turbidity. This could lead to significant changes in the trophic interactions in the food web of lakes. Most fish use their vision for predation and the location of prey can be highly influenced by light level and clarity of the water environment. Turbidity is an optical property of water that causes light to be scattered and absorbed by particles and molecules. Turbidity is highly variable in lakes, due to seasonal changes in suspended sediments, algal blooms and wind-driven suspension of sediments especially in shallow waters. There is evidence that human activity has increased erosion leading to increased turbidity in aquatic systems. Turbidity could also play a significant role in distribution of fish. Turbidity could act as a cover for small fish and reduce predation risk. Diel horizontal migration by fish is common in shallow lakes and is considered as consequences of either optimal foraging behaviour for food or as a trade-off between foraging and predator avoidance. In turbid lakes, diel horizontal migration patterns could differ since turbidity can act as a refuge itself and affect the predator-prey interactions. Laboratory experiments were conducted with perch (Perca fluviatilis L.) and white bream (Abramis björkna (L.)) to clarify the effects of turbidity on their feeding. Additionally to clarify the effects of turbidity on predator preying on different types of prey, pikeperch larvae (Sander lucioperca (L.)), Daphnia pulex (Leydig), Sida crystallina (O.F. Müller), and Chaoborus flavicans (Meigen) were used as prey in different experiments. To clarify the role of turbidity in distribution and diel horizontal migration of perch, roach (Rutilus rutilus (L.)) and white bream, field studies were conducted in shallow turbid lakes. A clear and a turbid shallow lake were compared to investigate distribution of perch and roach in these two lakes in a 15-year study period. Feeding efficiency of perch and white bream was not significantly affected with increasing clay turbidity up to 50 NTU. The perch experiments with pikeperch larvae suggested that clay turbidity could act as a refuge especially at turbidity levels higher than 50 NTU. Perch experiments with different prey types suggested that pikeperch larvae probably use turbidity as a refuge better compared to Daphnia. Increase in turbidity probably has stronger affect on perch predating on plant-attached prey. The main findings of the thesis show that turbidity can play a significant role in distribution of fish. Perch and roach could use turbidity as refuge when macrophytes disappear while small perch may also use high turbidity as refuge when macrophytes are present. Floating-leaved macrophytes are probably good refuges for small fish in clay-turbid lakes and provide a certain level of turbidity and not too complex structure for refuge. The results give light to the predator-prey interactions in turbid environments. Turbidity of water should be taken in to account when studying the diel horizontal migrations and distribution of fish in shallow lakes.

Acoustic emission source location in composite structure by Voronoi construction using geodesic curve evolution

Conventional analytical/numerical methods employing triangulation technique are suitable for locating acoustic emission (AE) source in a planar structure without structural discontinuities. But these methods cannot be extended to structures with complicated geometry, and, also, the problem gets compounded if the material of the structure is anisotropic warranting complex analytical velocity models. A geodesic approach using Voronoi construction is proposed in this work to locate the AE source in a composite structure. The approach is based on the fact that the wave takes minimum energy path to travel from the source to any other point in the connected domain. The geodesics are computed on the meshed surface of the structure using graph theory based on Dijkstra's algorithm. By propagating the waves in reverse virtually from these sensors along the geodesic path and by locating the first intersection point of these waves, one can get the AE source location. In this work, the geodesic approach is shown more suitable for a practicable source location solution in a composite structure with arbitrary surface containing finite discontinuities. Experiments have been conducted on composite plate specimens of simple and complex geometry to validate this method.

Microstructural evolution of tungsten oxide thin films

Tungsten oxide thin films are of great interest due to their promising applications in various optoelectronic thin film devices. We have investigated the microstructural evolution of tungsten oxide thin films grown by DC magnetron sputtering on silicon substrate. The structural characterization and surface morphology were carried out using X-ray diffraction and Scanning Electron Microscopy (SEM). The as deposited films were amorphous, where as, thin films annealed above 400 degrees C were crystalline. In order to explain the microstructural changes due to annealing, we have proposed a ``instability wheel'' model for the evolution of the microstructure. This model explains the transformation of mater into various geometries within them selves, followed by external perturbation.

Phylogeny of a taxonomically difficult group and evolution of host location mechanism

The parasitic wasps are one of the largest insect groups and their life histories are remarkably variable. Common to all parasitic wasps is that they kill their hosts, which are usually beetles, butterflies and sometimes spiders. Hosts are often at a larval or pupal stage and live in concealed conditions, such as in plant tissue. Parasitic wasps have two main ways of finding their host. 1) They can detect chemical compounds emitted by damaged plant material or released by larvae living in plant tissue, and 2) detect the larvae by sound vibrations. Even though pupae are immobile and silent, and therefore do not cause vibration, parasitoids have, however, adapted to find passive developmental stages by producing vibration themselves by knocking the substrate with their antennae, and then detecting the echoes with their legs. This echolocation allows a parasitoid to locate its potential hosts that are deeply buried in wood. This study focuses on the relationships of the subfamily Cryptinae (Hymenoptera: Ichneumonidae) and related taxa, and the evolution of host location mechanism. There are no earlier studies of the phylogeny of the Cryptinae, and the position of related taxa are unclear. According to the earlier classification, which is entirely intuitional, the Cryptinae is divided into three tribes: Cryptini, Hemigasterini and Phygadeuontini. Further, these tribes are subdiveded into numerous subtribes. This work, based on molecular characters, shows that the cryptine tribes Cryptini, Phygadeuon¬tini and Hemigasterini come out largely as monophyletic groups, thus agreeing with the earlier classification. The earlier subtribal classification had no support. In addition, it is shown that modified antennal structures are associated with host usage of wood-boring coleopteran hosts. The cryptines have a clear modification series on their antennal tips from a simply tip to a hammer-like structure. The species with strongly modified antennae belong mostly to the tribe Cryptini and they utilise wood-boring beetles as hosts. Also, field observations on insect behaviour support this result.

Oh Holly, The Fish is Dead

From Kurt Vonnegut to Stephen King, many novelists use metanarrative techniques to insert fictional versions of themselves in the stories they tell. The function of deploying such techniques is often to draw attention to the liminal space between the fictional constructs inherent in the novel as a form, and the real world from which the constructs draw inspiration, and indeed, are read by an audience. For emerging writers working in short form narratives, however, the structural demands of the short story or flash fiction make the use of similar techniques problematic in the level of depth to which they can be deployed. ‘Oh Holly, the fish is dead’ is the fourth in a series of short stories that work to overcome the structural limitations of a succinct form by developing a fractured fictional version of the author over a number of pieces and published across a range of sites. The accumulative affect is a richer metanarrative textual arrangement that also allows for the individual short stories to function independently.

lac Repressor Is an Antivirulence Factor of Salmonella enterica: Its Role in the Evolution of Virulence in Salmonella

The genus Salmonella includes many pathogens of great medical and veterinary importance. Bacteria belonging to this genus are very closely related to those belonging to the genus Escherichia. lacZYA operon and lacI are present in Escherichia coli, but not in Salmonella enterica. It has been proposed that Salmonella has lost lacZYA operon and lacI during evolution. In this study, we have investigated the physiological and evolutionary significance of the absence of lacI in Salmonella enterica. Using murine model of typhoid fever, we show that the expression of Lacl causes a remarkable reduction in the virulence of Salmonella enterica. Lacl also suppresses the ability of Salmonella enterica to proliferate inside murine macrophages. Microarray analysis revealed that Lacl interferes with the expression of virulence genes of Salmonella pathogenicity island 2. This effect was confirmed by RT-PCR and Western blot analysis. Interestingly, we found that SBG0326 of Salmonella bongori is homologous to lacI of Escherichia coli. Salmonella bongori is the only other species of the genus Salmonella and it lacks the virulence genes of Salmonella pathogenicity island 2. Overall, our results demonstrate that Lacl is an antivirulence factor of Salmonella enterica and suggest that absence of lacI has facilitated the acquisition of virulence genes of Salmonella pathogenicity island 2 in Salmonella enterica making it a successful systemic pathogen.

Third post-Newtonian angular momentum flux and the secular evolution of orbital elements for inspiralling compact binaries in quasi-elliptical orbits

The angular-momentum flux from an inspiralling binary system of compact objects moving in quasi-elliptical orbits is computed at the third post-Newtonian (3PN) order using the multipolar post-Minkowskian wave generation formalism. The 3PN angular-momentum flux involves the instantaneous, tail, and tail-of-tails contributions as for the 3PN energy flux, and in addition a contribution due to nonlinear memory. We average the angular-momentum flux over the binary's orbit using the 3PN quasi-Keplerian representation of elliptical orbits. The averaged angular-momentum flux provides the final input needed for gravitational-wave phasing of binaries moving in quasi-elliptical orbits. We obtain the evolution of orbital elements under 3PN gravitational radiation reaction in the quasi-elliptic case. For small eccentricities, we give simpler limiting expressions relevant for phasing up to order e(2). This work is important for the construction of templates for quasi-eccentric binaries, and for the comparison of post-Newtonian results with the numerical relativity simulations of the plunge and merger of eccentric binaries.

The evolution of loop structures in flux rings within the solar convection zone

Choudhuri and Gilman (1987) considered certain implications of the hypothesis that the magnetic flux within the Sun is generated at the bottom of the convection zone and then rises through it. Taking flux rings symmetric around the rotation axis and using reasonable values of different parameters, they found that the Coriolis force deflects these flux rings into trajectories parallel to the rotation axis so that they emerge at rather high latitudes. This paper looks into the question of whether the action of the Coriolis force is subdued when the initial configuration of the flux ring has non-axisymmetries in the form of loop structures. The results depend dramatically on whether the flux ring with the loops lies completely within the convection zone or whether the lower parts of it are embedded in the stable layers underneath the convection zone. In the first case, the Coriolis force supresses the non-axisymmetric perturbations so that the flux ring tends to remain symmetric and the trajectories are very similar to those of Choudhuri and Gilman (1987). In the second case, however, the lower parts of the flux ring may remain anchored underneath the bottom of the convection zone, but the upper parts of the loops still tend to move parallel to the rotation axis and emerge at high latitudes. Thus the problem of the magnetic flux not being able to come out at the sunspot latitudes still persists after the non-axisymmetries in the flux rings are taken into account.

Fish-scale bio-inspired multifunctional ZnO nanostructures

Scales provide optical disguise, low water drag and mechanical protection to fish, enabling them to survive catastrophic environmental disasters, predators and microorganisms. The unique structures and stacking sequences of fish scales inspired the fabrication of artificial nanostructures with salient optical, interfacial and mechanical properties. Herein, we describe fish-scale bio-inspired multifunctional ZnO nanostructures that have similar morphology and structure to the cycloid scales of the Asian Arowana. These nanostructured coatings feature tunable light refraction and reflection, modulated surface wettability and damage-tolerant mechanical properties. The salient properties of these multifunctional nanostructures are promising for applications in: - (i) optical coatings, sensing or lens arrays for use in reflective displays, packing, advertising and solar energy harvesting; - (ii) self-cleaning surfaces, including anti-smudge, anti-fouling and anti-fogging, and self-sterilizing surfaces, and; - (iii) mechanical/chemical barrier coatings. This study provides a low-cost and large-scale production method for the facile fabrication of these bio-inspired nanostructures and provides new insights for the development of novel functional materials for use in 'smart' structures and applications.