Vivek Nagendra Prakash

A bilateral body plan is predominant throughout the animal kingdom. Bilaterality of amniote embryos becomes recognizable as midline morphogenesis begins at gastrulation, bisecting an embryonic field into the left and right sides, and left-right (LR) asymmetry patterning follows. While a series of laterality genes expressed after the LR compartmentalization has been extensively studied, the laterality patterning prior to and at the initiation of midline morphogenesis has remained unclear. Here, through a biophysical quantification in a high spatial and temporal resolution, applied to a chick model system, we show that a large-scale bilateral counterrotating cellular flow, termed "polonaise movements", display LR asymmetries in early gastrulation. This cell movement starts prior to the formation of the primitive streak (PS) (the earliest midline structure) and the subsequent appearance of Hensen's node (the LR organizer). The cellular flow speed and vorticity unravel the location and timing of the LR asymmetries. The bilateral flows displayed a Right dominance after 6 h since the start of cell movements. Mitotic arrest that diminishes PS formation resulted in changes in the bilateral flow pattern, but the Right dominance persisted. Our data indicate that the LR asymmetry in amniote gastrula becomes detectable earlier than suggested by current models, which assume that the asymmetric regulation of the laterality signals at the node leads to the LR patterning. More broadly, our results suggest that physical processes can play an unexpected but significant role in influencing LR laterality during embryonic development.

In animals, epithelial tissues are barriers against the external environment, providing protection against biological, chemical, and physical damage. Depending on the organism's physiology and behavior, these tissues encounter different types of mechanical forces and need to provide a suitable adaptive response to ensure success. Therefore, understanding tissue mechanics in different contexts is an important research area. Here, we review recent tissue mechanics discoveries in three early-divergent non-bilaterian systems - Trichoplax adhaerens, Hydra vulgaris, and Aurelia aurita. We highlight each animal's simple body plan and biology, and unique, rapid tissue remodeling phenomena that play a crucial role in its physiology. We also discuss the emergent large-scale mechanics in these systems that arise from small-scale phenomena. Finally, we emphasize the potential of these non-bilaterian animals to be model systems in a bottom-up approach for further investigation in tissue mechanics.

ABSTRACT Most motile cilia have a stereotyped structure of nine microtubule outer doublets and a single central pair of microtubules. The central pair of microtubules are surrounded by a set of proteins, termed the central pair apparatus. A specific kinesin, Klp1 projects from the central pair and contributes to ciliary motility in Chlamydomonas. The vertebrate ortholog, Kif9, is required for beating in mouse sperm flagella, but the mechanism of Kif9/Klp1 function remains poorly defined. Here, using Xenopus epidermal multiciliated cells, we show that Kif9 is necessary for ciliary motility and the proper distal localization of not only central pair proteins, but also radial spokes and dynein arms. In addition, single-molecule assays in vitro reveal that Xenopus Kif9 is a long-range processive motor, although it does not mediate long-range movement in ciliary axonemes in vivo. Together, our data suggest that Kif9 is integral for ciliary beating and is necessary for proper axonemal distal end integrity.

Characterizing the epithelial tissue of a shape-shifting marine animal as an integrated composite material reveals a ductile-to-brittle phase transition that captures how the tissue responds to failure. Epithelial tissues provide an important barrier function in animals, but these tissues are subjected to extreme strains during day-to-day activities such as feeding and locomotion. Understanding tissue mechanics and the adaptive response in dynamic force landscapes remains an important area of research. Here we carry out a multi-modal study of a simple yet highly dynamic organism, Trichoplax adhaerens, and report the discovery of abrupt, bulk epithelial tissue fractures induced by the organism's own motility. Coupled with rapid healing, this discovery accounts for dramatic shape change and physiological asexual division in this early-divergent metazoan. We generalize our understanding of this phenomenon by codifying it in a heuristic model focusing on the debonding-bonding criterion in a soft, active living material. Using a suite of quantitative experimental and numerical techniques, we demonstrate a force-driven ductile-to-brittle material transition governing the morphodynamics of tissues pushed to the edge of rupture.

We present a simple, intuitive algorithm for visualizing time-varying flow fields that can reveal complex flow structures with minimal user intervention. We apply this technique to a variety of biological systems, including the swimming currents of invertebrates and the collective motion of swarms of insects. We compare our results with more experimentally difficult and mathematically sophisticated techniques for identifying patterns in fluid flows, and suggest that our tool represents an essential 'middle ground' allowing experimentalists to easily determine whether a system exhibits interesting flow patterns and coherent structures without resorting to more intensive techniques. In addition to being informative, the visualizations generated by our tool are often striking and elegant, illustrating coherent structures directly from videos without the need for computational overlays. Our tool is available as fully documented open-source code for MATLAB, Python or ImageJ at www.flowtrace.org.

Many marine invertebrates have larval stages covered in linear arrays of beating cilia, which propel the animal while simultaneously entraining planktonic prey(1). These bands are strongly conserved across taxa spanning four major superphyla(2,3), and they are responsible for the unusual morphologies of many invertebrate larvae(4,5). However, few studies have investigated their underlying hydrodynamics(6,7). Here, we study the ciliary bands of starfish larvae, and discover a beautiful pattern of slowly evolving vortices that surrounds the swimming animals. Closer inspection of the bands reveals unusual ciliary 'tangles' analogous to topological defects that break up and re-form as the animal adjusts its swimming stroke. Quantitative experiments and modelling demonstrate that these vortices create a physical trade-off between feeding and swimming in heterogeneous environments, which manifests as distinct flow patterns or 'eigenstrokes' representing each behaviour-potentially implicating neuronal control of cilia. This quantitative interplay between larval form and hydrodynamic function may generalize to other invertebrates with ciliary bands, and illustrates the potential effects of active boundary conditions in other biological and synthetic systems.