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

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

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

Ship and satellite bio-optical research in the California Bight

Mesoscale biological patterns and processes in productive coastal waters were studied. The physical and biological processes leading to chlorophyll variability were investigated. The ecological and evolutionary significance of this variability, and its relation to the prediction of fish recruitment and marine mammal distributions was studied. Seasonal primary productivity (using chlorophyll as an indication of phytoplankton biomass) for the entire Southern California Bight region was assessed. Complementary and contemporaneous ship and satellite (Nimbus 7-CZCS) bio-optical data from the Southern California Bight and surrounding waters were obtained and analyzed. These data were also utilized for the development of multi-platform sampling strategies and the optimization of algorithms for the estimation of phytoplankton biomass and primary production from satellite imagery

A mathematical basis for the clock and wavefront model for somitogenesis

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Distinguishing graded & ultrasensitive signalling cascade kinetics by the shape of morphogen gradients in Drosophila

Recently, signalling gradients in cascades of two-state reaction–diffusion systems were described as a model for understanding key biochemical mechanisms that underlie development and differentiation processes in the Drosophila embryo. Diffusion-trapping at the exterior of the cell membrane triggers the mitogen-activated protein kinase (MAPK) cascade to relay an appropriate signal from the membrane to the inner part of the cytosol, whereupon another diffusion-trapping mechanism involving the nucleus reads out this signal to trigger appropriate changes in gene expression. Proposed mathematical models exhibit equilibrium distributions consistent with experimental measurements of key spatial gradients in these processes. A significant property of the formulation is that the signal is assumed to be relayed from one system to the next in a linear fashion. However, the MAPK cascade often exhibits nonlinear dose–response properties and the final remark of Berezhkovskii et al. (2009) is that this assumption remains an important property to be tested experimentally, perhaps via a new quantitative assay across multiple genetic backgrounds. In anticipation of the need to be able to sensibly interpret data from such experiments, here we provide a complementary analysis that recovers existing formulae as a special case but is also capable of handling nonlinear functional forms. Predictions of linear and nonlinear signal relays and, in particular, graded and ultrasensitive MAPK kinetics, are compared

Laser anemometer measurements of trailing vortices in water

A series of measurements of trailing vortices behind lifting hydrofoils is described. These measurements were made in the Caltech Free-Surface Water Tunnel, using a laser-Doppler velocimeter to measure two components of velocity in the vortex wake. Two different model planforms were tested, and measurements were made at several free-stream velocities and angles of attack for each. Velocity profiles were measured at distances downstream of the model of from five to sixty chord lengths. These measurements are the first results of a continuing experimental programme. In § 3 of this paper, the theory of trailing vortices is discussed. The effects of ‘vortex wandering’ upon the measurements are computed, and the corrected results are seen to be in reasonable agreement with the theory