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Darwin’s finches are a classic example of adaptive radiation, exemplified by their adaptive and functional beak morphologies. To quantify their form, we carry out a morphometric analysis of the three-dimensional beak shapes of all of Darwin’s finches and find that they can be fit by a transverse parabolic shape with a curvature that increases linearly from the base toward the tip of the beak. The morphological variation of beak orientation, aspect ratios, and curvatures allows us to quantify beak function in terms of the elementary theory of machines, consistent with the dietary variations across finches. Finally, to explain the origin of the evolutionary morphometry and the developmental morphogenesis of the finch beak, we propose an experimentally motivated growth law at the cellular level that simplifies to a variant of curvature-driven flow at the tissue level and captures the range of observed beak shapes in terms of a simple morphospace. Altogether, our study illuminates how a minimal combination of geometry and dynamics allows for functional form to develop and evolve.

Bacterial growth is remarkably robust to environmental fluctuations, yet the mechanisms of growth-rate homeostasis are poorly understood. Here, we combine theory and experiment to infer mechanisms by which Escherichia coli adapts its growth rate in response to changes in osmolarity, a fundamental physicochemical property of the environment. The central tenet of our theoretical model is that cell-envelope expansion is only sensitive to local information, such as enzyme concentrations, cell-envelope curvature, and mechanical strain in the envelope. We constrained this model with quantitative measurements of the dynamics of E. coli elongation rate and cell width after hyperosmotic shock. Our analysis demonstrated that adaptive cell-envelope softening is a key process underlying growth-rate homeostasis. Furthermore, our model correctly predicted that softening does not occur above a critical hyperosmotic shock magnitude and precisely recapitulated the elongation-rate dynamics in response to shocks with magnitude larger than this threshold. Finally, we found that, to coordinately achieve growth-rate and cell-width homeostasis, cells employ direct feedback between cellenvelope curvature and envelope expansion. In sum, our analysis points to cellular mechanisms of bacterial growth-rate homeostasis and provides a practical theoretical framework for understanding this process.

Organs have a distinctive yet often overlooked spatial arrangement in the body 1 – 5 . We propose that there is a logic to the shape of an organ and its proximity to its neighbours. Here, by using volumetric scans of many Drosophila melanogaster flies, we develop methods to quantify three-dimensional features of organ shape, position and interindividual variability. We find that both the shapes of organs and their relative arrangement are consistent yet differ between the sexes, and identify unexpected interorgan adjacencies and left–right organ asymmetries. Focusing on the intestine, which traverses the entire body, we investigate how sex differences in three-dimensional organ geometry arise. The configuration of the adult intestine is only partially determined by physical constraints imposed by adjacent organs; its sex-specific shape is actively maintained by mechanochemical crosstalk between gut muscles and vascular-like trachea. Indeed, sex-biased expression of a muscle-derived fibroblast growth factor-like ligand renders trachea sexually dimorphic. In turn, tracheal branches hold gut loops together into a male or female shape, with physiological consequences. Interorgan geometry represents a previously unrecognized level of biological complexity which might enable or confine communication across organs and could help explain sex or species differences in organ function. In fruit flies, three-dimensional organ arrangement is stereotypical, sexually dimorphic and actively maintained by muscle-vessel mechanochemical crosstalk.

Thin shells are characterized by a high cost of stretching compared to bending. As a result isometries of the midsurface of a shell play a crucial role in their mechanics. In turn, curves with zero normal curvature play a critical role in determining the number and behavior of isometries. In this paper, we show how the presence of these curves results in a decrease in the number of linear isometries. Paradoxically, shells are also known to continuously fold more easily across these rigidifying curves than other curves on the surface. We show how including nonlinearities in the strain can explain this phenomena and demonstrate folding isometries with explicit solutions to the nonlinear isometry equations. In addition to explicit solutions, exact geometric arguments are given to validate and guide our analysis in a coordinate free way.

Adaptive avian radiations associated with the diversification of bird beaks into a multitude of forms enabling different functions are exemplified by Darwin's finches and Hawaiian honeycreepers. To elucidate the nature of these radiations, we quantified beak shape and skull shape using a variety of geometric measures that allowed us to collapse the variability of beak shape into a minimal set of geometric parameters. Furthermore, we find that just two measures of beak shape, the ratio of the width to length and the normalized sharpening rate (increase in the transverse beak curvature near the tip relative to that at the base of the beak), are both strongly correlated with diet, and thus indicative of how bite forces are correlated with beak shape. Finally, by considering how transverse sections to the beak centerline evolve with distance from the tip, we show that a simple geometry-driven growth law termed \"modified mean curvature flow\" captures the beak shapes of Darwin's finches and Hawaiian honeycreepers. A surprising consequence of the simple growth law is that beak shapes that are not allowed based on the developmental program of the beak are also not observed in nature, suggesting a link between evolutionary morphology and development in terms of growth-driven developmental constraints.Competing Interest StatementThe authors have declared no competing interest.

Safe, all-solid-state lithium metal batteries enable high energy density applications, but suffer from instabilities during operation that lead to rough interfaces between the metal and electrolyte and subsequently cause void formation and dendrite growth that degrades performance and safety. Inspired by the morphogenetic control of thin lamina such as tree leaves that robustly grow into flat shapes -- we propose a range of approaches to control lithium metal stripping and plating. To guide discovery of materials that will implement these feedback mechanisms, we develop a reduced order model that captures couplings between mechanics, interface growth, temperature, and electrochemical variables. We find that long-range feedback is required to achieve true interface stability, while approaches based on local feedback always eventually grow into rough interfaces. All together, our study provides the beginning of a practical framework for analyzing and designing stable electrochemical interfaces in terms of the mechanical properties and the physical chemistry that underlie their dynamics.

Temporal imaging of biological epithelial structures yields shape data at discrete time points, leading to a natural question: how can we reconstruct the most likely path of growth patterns consistent with these discrete observations? We present a physically plausible framework to solve this inverse problem by creating a framework that generalises quasiconformal maps to quasiconformal flows. By allowing for the spatio-temporal variation of the shear and dilatation fields during the growth process, subject to regulatory mechanisms, we are led to a type of generalised Ricci flow. When guided by observational data associated with surface shape as a function of time, this leads to a constrained optimization problem. Deploying our data-driven algorithmic approach to the shape of insect wings, leaves and even sculpted faces, we show how optimal quasiconformal flows allow us to characterise the morphogenesis of a range of surfaces.

Temporal imaging of biological epithelial structures yields shape data at discrete time points, leading to a natural question: how can we reconstruct the most likely path of growth patterns consistent with these discrete observations? We present a physically plausible framework to solve this inverse problem by creating a framework that generalises quasiconformal maps to quasiconformal flows. By allowing for the spatio-temporal variation of the shear and dilatation fields during the growth process, subject to regulatory mechanisms, we are led to a type of generalised Ricci flow. When guided by observational data associated with surface shape as a function of time, this leads to a constrained optimization problem. Deploying our data-driven algorithmic approach to the shape of insect wings, leaves and even sculpted faces, we show how optimal quasiconformal flows allow us to characterise the morphogenesis of a range of surfaces.

When a thin stream of aqueous sodium alginate is extruded into a reacting calcium chloride bath, it polymerizes into a soft elastic tube that spontaneously forms helical coils due to the ambient fluid drag. We quantify the onset of this drag-induced instability and its nonlinear evolution using experiments, and explain the results using a combination of scaling, theory and simulations. By co-extruding a second (internal) liquid within the aqueous sodium alginate jet and varying the rates of co-extrusion of the two liquids, as well as the diameter of the jet, we show that we can tune the local composition of the composite filament and the nature of the ensuing instabilities to create soft filaments of variable relative buoyancy, shape and mechanical properties. All together, by harnessing the fundamental varicose (jetting) and sinuous (buckling) instabilities associated with the extrusion of a jelling filament, we show that it is possible to print complex three-dimensional filamentous structures in the ambient fluid.

Bacterial growth is remarkably robust to environmental fluctuations, yet the mechanisms of growth-rate homeostasis are poorly understood. Here, we combine theory and experiment to infer mechanisms by which Escherichia coli adapts its growth rate in response to changes in osmolarity, a fundamental physicochemical property of the environment. The central tenet of our theoretical model is that cell-envelope expansion is only sensitive to local information such as enzyme concentrations, cell-envelope curvature, and mechanical strain in the envelope. We constrained this model with quantitative measurements of the dynamics of E. coli elongation rate and cell width after hyperosmotic shock. Our analysis demonstrated that adaptive cell-envelope softening is a key process underlying growth-rate homeostasis. Furthermore, our model correctly predicted that softening does not occur above a critical hyperosmotic shock magnitude and precisely recapitulated the elongation-rate dynamics in response to shocks with magnitude larger than this threshold. Finally, we found that to coordinately achieve growth-rate and cell-width homeostasis, cells employ direct feedback between cell-envelope curvature and envelope expansion. In sum, our analysis points to new cellular mechanisms of bacterial growth-rate homeostasis and provides a practical theoretical framework for understanding this process. The bacterial cell envelope is the critical structure that defines cell size and shape, and its expansion therefore defines cell growth. Although size, shape, and growth rate are important cellular variables that are robust to environmental fluctuations, the feedback mechanisms by which these variables influence cell-envelope expansion are unknown. Here, we explore how E. coli cells achieve growth-rate and cell-width homeostasis during fluctuations in osmolarity, a key environmental property. A biophysical model in which the cell envelope softens after an osmotic shock and envelope expansion depends directly on local curvature quantitatively recapitulated all experimental observations. Our study elucidates new mechanisms of bacterial cell morphogenesis and highlights the deep interplay between global cellular variables and the mechanisms of cell-envelope expansion.