Travelling gradients in interacting morphogen systems

Morphogen gradients are well known to play several important roles in development; however the mechanisms underlying the formation and maintenance of these gradients are often not well understood. In this work, we investigate whether the presence of a secondary morphogen can increase the robustness of the primary morphogen gradient to perturbation, thereby providing a more stable mechanism for development. We base our model around the interactions of Fibroblast Growth Factor 8 and retinoic acid, which have been shown to act as morphogens in many developmental systems. In particular, we investigate the formation of opposing gradients of these morphogens along the antero-posterior axis of vertebrate embryos, thereby controlling temporal and spatial aspects of axis segmentation and neuronal differentiation

Multiyear measurements of ebullitive methane flux from three subarctic lakes

Ebullition (bubbling) from small lakes and ponds at high latitudes is an important yet unconstrained source of atmospheric methane (CH4). Small water bodies are most abundant in permanently frozen peatlands, and it is speculated that their emissions will increase as the permafrost thaws. We made 6806 measurements of CH4 ebullition during four consecutive summers using a total of 40 bubble traps that were systematically distributed across the depth zones of three lakes in a sporadic permafrost landscape in northernmost Sweden. We identified significant spatial and temporal variations in ebullition and observed a large spread in the bubbles\u27 CH4 concentration, ranging from 0.04% to 98.6%. Ebullition followed lake temperatures, and releases were significantly larger during periods with decreasing atmospheric pressure. Although shallow zone ebullition dominated the seasonal bubble CH4 flux, we found a shift in the depth dependency towards higher fluxes from intermediate and deep zones in early fall. The average daily flux of 13.4 mg CH4 m−2 was lower than those measured in most other high‐latitude lakes. Locally, however, our study lakes are a substantial CH4 source; we estimate that 350 kg of CH4 is released via ebullition during summer (June–September), which is approximately 40% of total whole year emissions from the nearby peatland. In order to capture the large variability and to accurately scale lake CH4 ebullition temporally and spatially, frequent measurements over long time periods are critical

A mathematical investigation of a clock and wavefront model for somitogenesis

Abstract Somites are transient blocks of cells that form sequentially along the antero-posterior axis of vertebrate embryos. They give rise to the vertebrae, ribs and other associated features of the trunk. In this work we develop and analyse a mathematical formulation of a version of the Clock and Wavefront model for somite formation, where the clock controls when the boundaries of the somites form and the wavefront determines where they form. Our analysis indicates that this interaction between a segmentation clock and a wavefront can explain the periodic pattern of somites observed in normal embryos. We can also show that a simplification of the model provides a mechanism for predicting the anomalies resulting from perturbation of the wavefront

Waves and patterning in developmental biology: vertebrate segmentation and feather bud formation as case studies

In this article we will discuss the integration of developmental patterning mechanisms with waves of competency that control the ability of a homogeneous field of cells to react to pattern forming cues and generate spatially heterogeneous patterns. We base our discussion around two well known patterning events that take place in the early embryo: somitogenesis and feather bud formation. We outline mathematical models to describe each patterning mechanism, present the results of numerical simulations and discuss the validity of each model in relation to our example patterning processes

Formation of vertebral precursors: Past models and future predictions

Disruption of normal vertebral development results from abnormal formation and segmentation of the vertebral precursors, called somites. Somitogenesis, the sequential formation of a periodic pattern along the antero-posterior axis of vertebrate embryos, is one of the most obvious examples of the segmental patterning processes that take place during embryogenesis and also one of the major unresolved events in developmental biology. We review the most popular models of somite formation: Cooke and Zeeman's clock and wavefront model, Meinhardt's reaction-diffusion model and the cell cycle model of Stern and co-workers, and discuss the consistency of each in the light of recent experimental findings concerning FGF-8 signalling in the presomitic mesoderm (PSM). We present an extension of the cell cycle model to take account of this new experimental evidence, which shows the existence of a determination front whose position in the PSM is controlled by FGF-8 signalling, and which controls the ability of cells to become competent to segment. We conclude that it is, at this stage, perhaps erroneous to favour one of these models over the others

Formation of vertebral precursors: Past models and future predictions

Disruption of normal vertebral development results from abnormal formation and segmentation of the vertebral precursors, called somites. Somitogenesis, the sequential formation of a periodic pattern along the antero-posterior axis of vertebrate embryos, is one of the most obvious examples of the segmental patterning processes that take place during embryogenesis and also one of the major unresolved events in developmental biology. We review the most popular models of somite formation: Cooke and Zeeman's clock and wavefront model, Meinhardt's reaction-diffusion model and the cell cycle model of Stern and co-workers, and discuss the consistency of each in the light of recent experimental findings concerning FGF-8 signalling in the presomitic mesoderm (PSM). We present an extension of the cell cycle model to take account of this new experimental evidence, which shows the existence of a determination front whose position in the PSM is controlled by FGF-8 signalling, and which controls the ability of cells to become competent to segment. We conclude that it is, at this stage, perhaps erroneous to favour one of these models over the others

Plasma Wakefield Acceleration with a Modulated Proton Bunch

The plasma wakefield amplitudes which could be achieved via the modulation of a long proton bunch are investigated. We find that in the limit of long bunches compared to the plasma wavelength, the strength of the accelerating fields is directly proportional to the number of particles in the drive bunch and inversely proportional to the square of the transverse bunch size. The scaling laws were tested and verified in detailed simulations using parameters of existing proton accelerators, and large electric fields were achieved, reaching 1 GV/m for LHC bunches. Energy gains for test electrons beyond 6 TeV were found in this case.Comment: 9 pages, 7 figure