Just as in clay moulding or glass blowing, physically sculpting biological structures requires the constituent material to locally flow like a fluid while maintaining overall mechanical integrity like a solid. Disordered soft materials, such as foams, emulsions and colloidal suspensions, switch from fluid-like to solid-like behaviours at a jamming transition. Similarly, cell collectives have been shown to display glassy dynamics in 2D and 3D and jamming in cultured epithelial monolayers, behaviours recently predicted theoretically and proposed to influence asthma pathobiology and tumour progression. However, little is known about whether these seemingly universal behaviours occur in vivo and, specifically, whether they play any functional part during embryonic morphogenesis. Here, by combining direct in vivo measurements of tissue mechanics with analysis of cellular dynamics, we show that during vertebrate body axis elongation, posterior tissues undergo a jamming transition from a fluid-like behaviour at the extending end, the mesodermal progenitor zone, to a solid-like behaviour in the presomitic mesoderm. We uncover an anteroposterior, N-cadherin-dependent gradient in yield stress that provides increasing mechanical integrity to the presomitic mesoderm, consistent with the tissue transiting from a wetter to a dryer foam-like architecture. Our results show that cell-scale stresses fluctuate rapidly (within about 1 min), enabling cell rearrangements and effectively 'melting' the tissue at the growing end. Persistent (more than 0.5 h) stresses at supracellular scales, rather than cell-scale stresses, guide morphogenetic flows in fluid-like tissue regions. Unidirectional axis extension is sustained by the reported rigidification of the presomitic mesoderm, which mechanically supports posterior, fluid-like tissues during remodelling before their maturation. The spatiotemporal control of fluid-like and solid-like tissue states may represent a generic physical mechanism of embryonic morphogenesis.
It is generally believed that the mechanical properties of the cellular microenvironment and their spatiotemporal variations play a central role in sculpting embryonic tissues, maintaining organ architecture and controlling cell behavior, including cell differentiation. However, no direct in vivo and in situ measurement of mechanical properties within developing 3D tissues and organs has been performed yet. Here we introduce a technique that employs biocompatible ferrofluid microdroplets as local mechanical actuators and allows quantitative spatiotemporal measurements of mechanical properties in vivo. Using this technique, we show that vertebrate body elongation entails spatially-varying tissue mechanics along the anteroposterior axis. Specifically, we find that the zebrafish tailbud is viscoelastic (elastic below a few seconds and fluid after just one minute) and displays decreasing stiffness and increasing fluidity towards its posterior elongating region. This method opens new avenues to study mechanobiology in vivo, both in embryogenesis and in disease processes, including cancer.
We consider DNA translocation through a pore in a planar membrane. The pore is so narrow that only one DNA segment can fit in. Assuming that the biasing force f acts inside the pore only, and that the DNA monomer number N is asymptotically large, we modify the previously developed treatment of the stretched part of the pre-translocated polymer by introducing the concept of "iso-flux trumpet". We show that friction of a moving chain in the trumpet, although it determines the speed of the process, provides only a marginal fraction of overall dissipation in the process. The dominant dissipation turns out to be due to irreversible entropic squeezing of the chain into the small pore. We also discover that because of the role of the membrane a much larger amount of heat of order k(B)T per monomer gets transferred from the heat bath on the post-translocation side to that on the pre-translocation side.
Based on our formulation of the DNA electrophoresis near a pore [Rowghanian and Grosberg, Phys. Rev. E (to be published)], we address the electrophoretic DNA capture into a nanopore as a steady-state process of particle absorption to a sink placed on top of an energy barrier. Reproducing the previously observed diffusion-limited and barrier-limited regimes as two different limits of the particle absorption process and matching the data, our model suggests a slower growth of the capture rate with the DNA length for very large DNA molecules than the previous model, motivating more experiments beyond the current range of electric field and DNA length. At moderately weak electric fields, our model predicts a different effect, stating that the DNA length dependence of the capture rate first disappears as the field is reduced and eventually reverses to a decreasing trend with N.
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