Deformations of cell sheets are ubiquitous in early animal development, often arising from a complex and poorly understood interplay of cell shape changes, division, and migration. Here, we explore perhaps the simplest example of cell sheet folding: the "inversion" process of the algal genus Volvox, during which spherical embryos turn themselves inside out through a process hypothesized to arise from cell shape changes alone. We use light sheet microscopy to obtain the first three-dimensional visualizations of inversion in vivo, and develop the first theory of this process, in which cell shape changes appear as local variations of intrinsic curvature, contraction and stretching of an elastic shell. Our results support a scenario in which these active processes function in a defined spatiotemporal manner to enable inversion.
Variability is emerging as an integral part of development. It is therefore imperative to ask how to access the information contained in this variability. Yet most studies of development average their observations and, discarding the variability, seek to derive models, biological or physical, that explain these average observations. Here, we analyse this variability in a study of cell sheet folding in the green alga Volvox, whose spherical embryos turn themselves inside out in a process sharing invagination, expansion, involution, and peeling of a cell sheet with animal models of morphogenesis. We generalise our earlier, qualitative model of the initial stages of inversion by combining ideas from morphoelasticity and shell theory. Together with three-dimensional visualisations of inversion using light sheet microscopy, this yields a detailed, quantitative model of the entire inversion process. With this model, we show how the variability of inversion reveals that two separate, temporally uncoupled processes drive the initial invagination and subsequent expansion of the cell sheet. This implies a prototypical transition towards higher developmental complexity in the volvocine algae and provides proof of principle of analysing morphogenesis based on its variability.
Recent studies of cooled oil emulsion droplets uncovered transformations into a host of flattened shapes with straight edges and sharp corners, driven by a partial phase transition of the bulk liquid phase. Here, we explore theoretically the simplest geometric competition between this phase transition and surface tension in planar polygons and recover the observed sequence of shapes and their statistics in qualitative agreement with experiments. Extending the model to capture some of the three-dimensional structure of the droplets, we analyze the evolution of protrusions sprouting from the vertices of the platelets and the topological transition of a puncturing planar polygon.
In microbial communities, each species may have multiple, distinct phenotypes. How do these subpopulations affect the stability of the community? Here, we address this question theoretically, showing that simple models with subpopulation structure and averaged versions thereof generically give contradictory linear stability results. Specialising to the bacterial persister phenotype, we analyse stochastic switching between phenotypes in detail in an asymptotic limit. Abundant phenotypic variation tends to be linearly destabilising, but, surprisingly, a rare phenotype can have a stabilising effect. Finally, we extend these results by showing numerically that subpopulations also modify the stability of the system to large perturbations such as antibiotic treatments.
Elastic objects across a wide range of scales deform under local changes of their intrinsic properties, yet the shapes are glocal, set by a complicated balance between local properties and global geometric constraints. Here, we explore this interplay during the inversion process of the green alga Volvox, whose embryos must turn themselves inside out to complete their development. This process has recently been shown to be well described by the deformations of an elastic shell under local variations of its intrinsic curvatures and stretches, although the detailed mechanics of the process have remained unclear. Through a combination of asymptotic analysis and numerical studies of the bifurcation behaviour, we illustrate how appropriate local deformations can overcome global constraints to initiate inversion.
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