Nonlinear elasticity and short-range mechanical coupling govern the rate and symmetry of mouth opening in Hydra.

Hydra has a tubular bilayered epithelial body column with a dome-shaped head on one end and a foot on the other. Hydra lacks a permanent mouth: its head epithelium is sealed. Upon neuronal activation, a mouth opens at the apex of the head which can exceed the body column diameter in seconds, allowing Hydra to ingest prey larger than itself. While the kinematics of mouth opening are well characterized, the underlying mechanism is unknown. We show that Hydra mouth opening is generated by independent local contractions that require tissue-level coordination. We model the head epithelium as an active viscoelastic nonlinear spring network. The model reproduces the size, timescale and symmetry of mouth opening. It shows that radial contractions, travelling inwards from the outer boundary of the head, pull the mouth open. Nonlinear elasticity makes mouth opening larger and faster, contrary to expectations. The model correctly predicts changes in mouth shape in response to external forces. By generating innervated : nerve-free chimera in experiments and simulations, we show that nearest-neighbour mechanical signalling suffices to coordinate mouth opening. Hydra mouth opening shows that in the absence of long-range chemical or neuronal signals, short-range mechanical coupling is sufficient to produce long-range order in tissue deformations.

How the birth and death of shear layers determine confinement evolution: from the L → H transition to the density limit.

Electric field profile structure-especially its shear-is a natural order parameter for the edge plasma, and characterizes confinement regimes ranging from the H-mode (Wagner et al. 1982 Phys. Rev. Lett. 49, 1408-1412 (doi:10.1103/PhysRevLett.49.1408)) to the density limit (DL) (Greenwald et al. 1988 Nucl. Fusion 28, 2199-2207 (doi:10.1088/0029-5515/28/12/009)). The theoretical developments and lessons learned during 40 years of H-mode studies (Connor & Wilson 1999 Plasma Phys. Control. Fusion 42, R1-R74 (doi:10.1088/0741-3335/42/1/201); Wagner 2007 Plasma Phys. Control. Fusion 49, B1-B33 (doi:10.1088/0741-3335/49/12b/s01)) are applied to the shear layer collapse paradigm (Hong et al. 2017 Nucl. Fusion 58, 016041 (doi:10.1088/1741-4326/aa9626)) for the onset of DL phenomena. Results from recent experiments on edge shear layers and DL phenomenology are summarized and discussed in the light of L [Formula: see text] H transition physics. The theory of shear layer collapse is then developed. We demonstrate that shear layer physics captures both the well known current (Greenwald) scaling of the DL (Greenwald 2002 Plasma Phys. Control. Fusion 44, R27-R53 (doi:10.1088/0741-3335/44/8/201); Greenwald et al. 2014 Phys. Plasmas 21, 110501 (doi:10.1063/1.4901920)), as well as the emerging power scaling (Zanca, Sattin, JET Contributors 2019 Nucl. Fusion 59, 126011 (doi:10.1088/1741-4326/ab3b31)). The derivation of the power scaling theory exploits an existing model, originally developed for the L [Formula: see text] H transition (Diamond, Liang, Carreras, Terry 1994 Phys. Rev. Lett. 72, 2565-2568 (doi:10.1103/PhysRevLett.72.2565); Kim & Diamond 2003 Phys. Rev. Lett. 90, 185006 (doi:10.1103/PhysRevLett.90.185006)). We describe the enhanced particle transport events that occur following shear layer collapse. Open problems and future directions are discussed. This article is part of a discussion meeting issue 'H-mode transition and pedestal studies in fusion plasmas'.

Let it rip: the mechanics of self-bisection in asexual planarians determines their population reproductive strategies.

Asexual freshwater planarians reproduce by transverse bisection (binary fission) into two pieces. This process produces a head and a tail, which fully regenerate within 1-2 weeks. How planarians split into two offspring-using only their musculature and substrate traction-is a challenging biomechanics problem. We found that three different species,Dugesia japonica,Girardia tigrinaandSchmidtea mediterranea, have evolved three different mechanical solutions to self-bisect. Using time lapse imaging of the fission process, we quantitatively characterize the main steps of division in the three species and extract the distinct and shared key features. Across the three species, planarians actively alter their body shape, regulate substrate traction, and use their muscles to generate tensile stresses large enough to overcome the ultimate tensile strength of the tissue. Moreover, we show thathoweach planarian species divides dictates how resources are split among its offspring. This ultimately determines offspring survival and reproductive success. Thus, heterospecific differences in the mechanics of self-bisection of individual worms explain the observed differences in the population reproductive strategies of different planarian species.

Mouth Function Determines the Shape Oscillation Pattern in Regenerating Hydra Tissue Spheres.

Hydra is a small freshwater polyp capable of regeneration from small tissue pieces and from aggregates of cells. During regeneration, a hollow bilayered sphere is formed that undergoes osmotically driven shape oscillations of inflation and rupture. These oscillations are necessary for successful regeneration. Eventually, the oscillating sphere breaks rotational symmetry along the future head-foot axis of the animal. Notably, the shape oscillations show an abrupt shift from large-amplitude, long-period oscillations to small-amplitude, short-period oscillations. It has been widely accepted that this shift in oscillation pattern is linked to symmetry breaking and axis formation, and current theoretical models of Hydra symmetry breaking use this assumption as a model constraint. However, a mechanistic explanation for the shift in oscillation pattern is lacking. Using in vivo manipulation and imaging, we quantified the shape oscillation dynamics and dissected the timing and triggers of the pattern shift. Our experiments demonstrate that the shift in the shape oscillation pattern in regenerating Hydra tissue pieces is caused by the formation of a functional mouth and not by shape symmetry breaking as previously assumed. Thus, model assumptions must be revised in light of these new experimental data, which can be used to constrain and validate improved theoretical models of pattern formation in Hydra.

Mechanics dictate where and how freshwater planarians fission.

Asexual freshwater planarians reproduce by tearing themselves into two pieces by a process called binary fission. The resulting head and tail pieces regenerate within about a week, forming two new worms. Understanding this process of ripping oneself into two parts poses a challenging biomechanical problem. Because planarians stop "doing it" at the slightest disturbance, this remained a centuries-old puzzle. We focus on Dugesia japonica fission and show that it proceeds in three stages: a local constriction ("waist formation"), pulsation-which increases waist longitudinal stresses-and transverse rupture. We developed a linear mechanical model with a planarian represented by a thin shell. The model fully captures the pulsation dynamics leading to rupture and reproduces empirical time scales and stresses. It asserts that fission execution is a mechanical process. Furthermore, we show that the location of waist formation, and thus fission, is determined by physical constraints. Together, our results demonstrate that where and how a planarian rips itself apart during asexual reproduction can be fully explained through biomechanics.

How electron two-stream instability drives cyclic Langmuir collapse and continuous coherent emission.

Continuous plasma coherent emission is maintained by repetitive Langmuir collapse driven by the nonlinear evolution of a strong electron two-stream instability. The Langmuir waves are modulated by solitary waves in the linear stage and electrostatic whistler waves in the nonlinear stage. Modulational instability leads to Langmuir collapse and electron heating that fills in cavitons. The high pressure is released via excitation of a short-wavelength ion acoustic mode that is damped by electrons and reexcites small-scale Langmuir waves; this process closes a feedback loop that maintains the continuous coherent emission.