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Biological materials, in addition to having remarkable physical properties, can also change shape and volume. These shape and volume changes allow organisms to form new tissue during growth and morphogenesis, as well as to repair and remodel old tissues. In addition shape or volume changes in an existing tissue can lead to useful motion or force generation (actuation) that may even still function in the dead organism, such as in the well known example of the hygroscopic opening or closing behaviour of the pine cone. Both growth and actuation of tissues are mediated, in addition to biochemical factors, by the physical constraints of the surrounding environment and the architecture of the underlying tissue. This habilitation thesis describes biophysical studies carried out over the past years on growth and swelling mediated shape changes in biological systems. These studies use a combination of theoretical and experimental tools to attempt to elucidate the physical mechanisms governing geometry controlled tissue growth and geometry constrained tissue swelling. It is hoped that in addition to helping understand fundamental processes of growth and morphogenesis, ideas stemming from such studies can also be used to design new materials for medicine and robotics.
Kaliumionen (K<sup>+) sind die am häufigsten vorkommenden anorganischen Kationen in Pflanzen. Gemessen am Trockengewicht kann ihr Anteil bis zu 10% ausmachen. Kaliumionen übernehmen wichtige Funktionen in verschiedenen Prozessen in der Pflanze. So sind sie z.B. essentiell für das Wachstum und für den Stoffwechsel. Viele wichtige Enzyme arbeiten optimal bei einer K<sup>+ Konzentration im Bereich von 100 mM. Aus diesem Grund halten Pflanzenzellen in ihren Kompartimenten, die am Stoffwechsel beteiligt sind, eine kontrollierte Kaliumkonzentration von etwa 100 mM aufrecht. Die Aufnahme von Kaliumionen aus dem Erdreich und deren Transport innerhalb der Pflanze und innerhalb einer Pflanzenzelle wird durch verschiedene Kaliumtransportproteine ermöglicht. Die Aufrechterhaltung einer stabilen K<sup>+ Konzentration ist jedoch nur möglich, wenn die Aktivität dieser Transportproteine einer strikten Kontrolle unterliegt. Die Prozesse, die die Transportproteine regulieren, sind bis heute nur ansatzweise verstanden. Detailliertere Kenntnisse auf diesem Gebiet sind aber von zentraler Bedeutung für das Verständnis der Integration der Transportproteine in das komplexe System des pflanzlichen Organismus. In dieser Habilitationsschrift werden eigene Publikationen zusammenfassend dargestellt, in denen die Untersuchungen verschiedener Regulationsmechanismen pflanzlicher Kaliumkanäle beschrieben werden. Diese Untersuchungen umfassen ein Spektrum aus verschiedenen proteinbiochemischen, biophysikalischen und pflanzenphysiologischen Analysen. Um die Regulationsmechanismen grundlegend zu verstehen, werden zum einen ihre strukturellen und molekularen Besonderheiten untersucht. Zum anderen werden die biophysikalischen und reaktionskinetischen Zusammenhänge der Regulationsmechanismen analysiert. Die gewonnenen Erkenntnisse erlauben eine neue, detailliertere Interpretation der physiologischen Rolle der Kaliumtransportproteine in der Pflanze.
The behaviour of an adhering cell is strongly influenced by the chemical, topographical and mechanical properties of the surface it attaches to. During recent years, it has been found experimentally that adhering cells actively sense the elastic properties of their environment by pulling on it through numerous sites of adhesion. The resulting build-up of force at sites of adhesion depends on the elastic properties of the environment and is converted into corresponding biochemical signals, which can trigger cellular programmes like growth, differentiation, apoptosis, and migration. In general, force is an important regulator of biological systems, for example in hearing and touch, in wound healing, and in rolling adhesion of leukocytes on vessel walls. In the habilitation thesis by Ulrich Schwarz, several theoretical projects are presented which address the role of forces and elasticity in cell adhesion. (1) A new method has been developed for calculating cellular forces exerted at sites of focal adhesion on micro-patterned elastic substrates. The main result is that cell-matrix contacts function as mechanosensors, converting internal force into protein aggregation. (2) A one-step master equation for the stochastic dynamics of adhesion clusters as a function of cluster size, rebinding rate and force has been solved both analytically and numerically. Moreover this model has been applied to the regulation of cell-matrix contacts, to dynamic force spectroscopy, and to rolling adhesion. (3) Using linear elasticity theory and the concept of force dipoles, a model has been introduced and solved which predicts the positioning and orientation of mechanically active cells in soft material, in good agreement with experimental observations for fibroblasts on elastic substrates and in collagen gels.