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A fundamental issue in biology is the nature of evolutionary transitions from unicellular to multicellular organisms. Volvocine algae are models for this transition, as they span from the unicellular biflagellate Chlamydomonas to multicellular species of Volvox with up to 50,000 Chlamydomonas-like cells on the surface of a spherical extracellular matrix. The mechanism of phototaxis in these species is of particular interest since they lack a nervous system and intercellular connections; steering is a consequence of the response of individual cells to light. Studies of Volvox and Gonium, a 16-cell organism with a plate-like structure, have shown that the flagellar response to changing illumination of the cellular photosensor is adaptive, with a recovery time tuned to the rotation period of the colony around its primary axis. Here, combining high-resolution studies of the flagellar photoresponse with 3D tracking of freely-swimming cells, we show that such tuning also underlies phototaxis of Chlamydomonas. A mathematical model is developed based on the rotations around an axis perpendicular to the flagellar beat plane that occur through the adaptive response to oscillating light levels as the organism spins. Exploiting a separation of time scales between the flagellar photoresponse and phototurning, we develop an equation of motion that accurately describes the observed photoalignment. In showing that the adaptive time scale is tuned to the organisms' rotational period across three orders of magnitude in cell number, our results suggest a unified picture of phototaxis in green algae in which the asymmetry in torques that produce phototurns arise from the individual flagella of Chlamydomonas, the flagellated edges of Gonium and the flagellated hemispheres of Volvox., Comment: 24 pages, 21 figures, 3 supplementary videos (available from REG)

The Rosetta spacecraft detected transient and sporadic diamagnetic regions around comet 67P/Churyumov-Gerasimenko. In this paper we present a statistical analysis of bulk and suprathermal electron dynamics, as well as a case study of suprathermal electron pitch angle distributions (PADs) near a diamagnetic region. Bulk electron densities are correlated with the local neutral density and we find a distinct enhancement in electron densities measured over the southern latitudes of the comet. Flux of suprathermal electrons with energies between tens of eV to a couple of hundred eV decreases each time the spacecraft enters a diamagnetic region. We propose a mechanism in which this reduction can be explained by solar wind electrons that are tied to the magnetic field and after having been transported adiabatically in a decaying magnetic field environment, have limited access to the diamagnetic regions. Our analysis shows that suprathermal electron PADs evolve from an almost isotropic outside the diamagnetic cavity to a field-aligned distribution near the boundary. Electron transport becomes chaotic and non-adiabatic when electron gyroradius becomes comparable to the size of the magnetic field line curvature, which determines the upper energy limit of the flux variation. This study is based on Rosetta observations at around 200 km cometocentric distance when the comet was at 1.24 AU from the Sun and during the southern summer cometary season.

We present observations from the Rosetta Plasma Consortium of the effects of stormy solar wind on comet 67P/Churyumov-Gerasimenko. Four corotating interaction regions (CIRs), where the first event has possibly merged with a CME, are traced from Earth via Mars (using Mars Express and MAVEN) and to comet 67P from October to December 2014. When the comet is 3.1-2.7 AU from the Sun and the neutral outgassing rate $\sim10^{25}-10^{26}$ s$^{-1}$ the CIRs significantly influence the cometary plasma environment at altitudes down to 10-30 km. The ionospheric low-energy \textcolor{black}{($\sim$5 eV) plasma density increases significantly in all events, by a factor $>2$ in events 1-2 but less in events 3-4. The spacecraft potential drops below -20V upon impact when the flux of electrons increases}. The increased density is \textcolor{black}{likely} caused by compression of the plasma environment, increased particle impact ionisation, and possibly charge exchange processes and acceleration of mass loaded plasma back to the comet ionosphere. During all events, the fluxes of suprathermal ($\sim$10-100 eV) electrons increase significantly, suggesting that the heating mechanism of these electrons is coupled to the solar wind energy input. At impact the magnetic field strength in the coma increases by a factor of ~2-5 as more interplanetary magnetic field piles up around of the comet. During two CIR impact events, we observe possible plasma boundaries forming, or moving past Rosetta, as the strong solar wind compresses the cometary plasma environment. \textcolor{black}{We also discuss the possibility of seeing some signatures of the ionospheric response to tail disconnection events, Comment: 17 pages, 8 figures

We present Rosetta observations from comet 67P/Churyumov-Gerasimenko during the impact of a coronal mass ejection (CME). The CME impacted on 5-6 Oct 2015, when Rosetta was about 800 km from the comet nucleus, \textcolor{black}{and 1.4 AU from the Sun}. Upon impact, the plasma environment is compressed to the level that solar wind ions, not seen a few days earlier when at 1500 km, now reach Rosetta. In response to the compression, the flux of suprathermal electrons increases by a factor of 5-10 and the background magnetic field strength increases by a factor of $\sim$2.5. The plasma density increases by a factor of 10 and reaches 600 cm$^{-3}$, due to increased particle impact ionisation, charge exchange and the adiabatic compression of the plasma environment. We also observe unprecedentedly large magnetic field spikes at 800 km, reaching above 200 nT, which are interpreted as magnetic flux ropes. We suggest that these could possibly be formed by magnetic reconnection processes in the coma as the magnetic field across the CME changes polarity, or as a consequence of strong shears causing Kelvin-Helmholtz instabilities in the plasma flow. Due to the \textcolor{black}{limited orbit of Rosetta}, we are not able to observe if a tail disconnection occurs during the CME impact, which could be expected based on previous remote observations of other CME-comet interactions., Comment: 14 pages, 12 figures

We use measurements from the Rosetta plasma consortium (RPC) Langmuir probe (LAP) and mutual impedance probe (MIP) to study the spatial distribution of low-energy plasma in the near-nucleus coma of comet 67P/Churyumov-Gerasimenko. The spatial distribution is highly structured with the highest density in the summer hemisphere and above the region connecting the two main lobes of the comet, i.e. the neck region. There is a clear correlation with the neutral density and the plasma to neutral density ratio is found to be about 1-2x10^-6, at a cometocentric distance of 10 km and at 3.1 AU from the sun. A clear 6.2 h modulation of the plasma is seen as the neck is exposed twice per rotation. The electron density of the collisonless plasma within 260 km from the nucleus falls of with radial distance as about 1/r. The spatial structure indicates that local ionization of neutral gas is the dominant source of low-energy plasma around the comet., Comment: 7 pages, 3 figures

Colonies of the green alga $Volvox$ are spheres that swim through the beating of pairs of flagella on their surface somatic cells. The somatic cells themselves are mounted rigidly in a polymeric extracellular matrix, fixing the orientation of the flagella so that they beat approximately in a meridional plane, with axis of symmetry in the swimming direction, but with a roughly 15 degree azimuthal offset which results in the eponymous rotation of the colonies about a body-fixed axis. Experiments on colonies held stationary on a micropipette show that the beating pattern takes the form of a symplectic metachronal wave (Brumley et al. (2012)). Here we extend the Lighthill/Blake axisymmetric, Stokes-flow model of a free-swimming spherical squirmer (Lighthill (1952), Blake (1971b)) to include azimuthal swirl. The measured kinematics of the metachronal wave for 60 different colonies are used to calculate the coefficients in the eigenfunction expansions and hence predict the mean swimming speeds and rotation rates, proportional to the square of the beating amplitude, as functions of colony radius. As a test of the squirmer model, the results are compared with measurements (Drescher et al. (2009)) of the mean swimming speeds and angular velocities of a different set of 220 colonies, also given as functions of colony radius. The predicted variation with radius is qualitatively correct, but the model underestimates both the mean swimming speed and the mean angular velocity unless the amplitude of the flagellar beat is taken to be larger than previously thought. The reasons for this discrepancy are discussed., Comment: 21 pages, 12 figures, 4 supplemental movies available on website of REG

We report on magnetic field measurements made in the innermost coma of 67P/Churyumov-Gerasimenko in its low activity state. Quasi-coherent, large-amplitude ($\delta B/B \sim 1$), compressional magnetic field oscillations at $\sim$ 40 mHz dominate the immediate plasma environment of the nucleus. This differs from previously studied comet-interaction regions where waves at the cometary ion gyro-frequencies are the main feature. Thus classical pick-up ion driven instabilities are unable to explain the observations. We propose a cross-field current instability associated with newborn cometary ion currents as a possible source mechanism., Comment: 6 pages, 3 Figures

Many structural properties of conventional passive materials are known to arise from the symmetries of their microscopic constituents. By contrast, it is largely unclear how the interplay between cell shape and self-propulsion controls the meso- and macroscale behavior of active matter. Here, we analyze large-scale simulations of homo- and heterogeneous self-propelled particle systems to identify generic effects of broken particle symmetry on collective motion. We find that even small violations of fore-aft symmetry lead to fundamentally different collective behaviors, which may facilitate demixing of differently shaped species as well as the spontaneous formation of stable micro-rotors. These results suggest that variation of particle shape yields robust physical mechanisms to control self-assembly of active matter, with possibly profound implications for biology and materials design., Comment: 5 pages, 4 figures

Intracellular cargo transport can arise from passive diffusion, active motor-driven transport along cytoskeletal filament networks, and passive advection by fluid flows entrained by such motor/cargo motion. Active and advective transport are thus intrinsically coupled as related, yet different representations of the same underlying network structure. A reaction-advection-diffusion system is used here to show that this coupling affects the transport and localization of a passive tracer in a confined geometry. For sufficiently low diffusion, cargo localization to a target zone is optimized either by low reaction kinetics and decoupling of bound and unbound states, or by a mostly disordered cytoskeletal network with only weak directional bias. These generic results may help to rationalize subtle features of cytoskeletal networks, for example as observed for microtubules in fly oocytes., Comment: revtex, 5 pages, 5 figures, to appear in PRL (http://prl.aps.org/)

Recently Bar-Ziv and Moses discovered a dynamical shape transformation induced in cylindrical lipid bilayer vesicles by the action of laser tweezers. We develop a hydrodynamic theory of fluid bilayers in interaction with the surrounding water and argue that the effect of the laser is to induce a sudden tension in the membrane. We refine our previous analysis to account for the fact that the shape transformation is not uniform but propagates outward from the laser trap. Applying the marginal stability criterion to this situation gives us an improved prediction for the selected initial wavelength and a new prediction for the propagation velocity, both in rough agreement with the experimental values. For example, a tubule of initial radius 0.7\micron\ has a predicted initial sinusoidal perturbation in its diameter with wavelength 5.5\micron, as observed. The perturbation propagates as a front with the qualitatively correct front velocity a bit less than 100\micron/sec. In particular we show why this velocity is initially constant, as observed, and so much smaller than the natural scale set by the tension. We also predict that the front velocity should increase linearly with laser power. Finally we introduce an approximate hydrodynamic model applicable to the fully nonlinear regime. This model exhibits propagating fronts as well as fully-developed ``pearled" vesicles similar to those seen in the experiments., Comment: 42 pages, 6 eps figures included with text in uuencoded file, ps file available from ftp://dept.physics.upenn.edu/pub/Nelson/pearl_propagation.ps submitted to Journal de Physique