Person: Berg, Howard
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Berg, Howard
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Publication Perspectives on working at the physics-biology interface(IOP Publishing, 2014) Berg, Howard; Blagoev, KrastanPublication Membrane Dipole Potentials(Elsevier BV, 1968) Berg, HowardPublication Determination of the Quadrupole Coupling Constant in the N^{14} Atomic Ground State(American Physical Society, 1970) Crampton, S. B.; Berg, Howard; Robinson, H. G.; Ramsey, NormanThe zero-field hyperfine frequencies in the ground state of atomic N14 have been measured in a hydrogen maser with sufficient accuracy to resolve the quadrupole coupling constant B. The result is B=+1.32±0.20 Hz.Publication Forced Axial Flow Between Rotating Concentric Cylinders(Cambridge University Press (CUP), 1971) Barcilon, Victor; Berg, HowardForced axial flow in an annular gap of a cylindrical rotor is investigated analytically and experimentally. At small rotation rates and narrow gap widths, the axial flow is a simple Poiseuille flow over most of the rotor. The distance required for this Poiseuille flow to get established is estimated. An instability is observed at large rotation rates with certain input geometries.Publication Cell motility: Turning failure into function(Springer Nature, 2013) Berg, HowardIn their search for more favourable environments bacteria choose new directions to explore, usually at random. In a marine bacterium with a single polar flagellum it is now shown that this quest is enhanced by a buckling instability.Publication Physics of chemoreception(Elsevier BV, 1977) Berg, Howard; Purcell, E.M.Statistical fluctuations limit the precision with which a microorganism can, in a given time T, determine the concentration of a chemoattractant in the surrounding medium. The best a cell can do is to monitor continually the state of occupation of receptors distributed over its surface. For nearly optimum performance only a small fraction of the surface need be specifically adsorbing. The probability that a molecule that has collided with the cell will find a receptor is Ns/(Ns + pi a), if N receptors, each with a binding site of radius s, are evenly distributed over a cell of radius a. There is ample room for many indenpendent systems of specific receptors. The adsorption rate for molecules of moderate size cannot be significantly enhanced by motion of the cell or by stirring of the medium by the cell. The least fractional error attainable in the determination of a concentration c is approximately (TcaD) - 1/2, where D is diffusion constant of the attractant. The number of specific receptors needed to attain such precision is about a/s. Data on bacteriophage absorption, bacterial chemotaxis, and chemotaxis in a cellular slime mold are evaluated. The chemotactic sensitivity of Escherichia coli approaches that of the cell of optimum design.Publication Adaptation Kinetics in Bacterial Chemotaxis(American Society for Microbiology, 1983) Block, Steven M.; Segall, Jeffrey E.; Berg, HowardCells of Escherichia coli, tethered to glass by a single flagellum, were subjected to constant flow of a medium containing the attractant alpha-methyl-DL-aspartate. The concentration of this chemical was varied with a programmable mixing apparatus over a range spanning the dissociation constant of the chemoreceptor at rates comparable to those experienced by cells swimming in spatial gradients. When an exponentially increasing ramp was turned on (a ramp that increases the chemoreceptor occupancy linearly), the rotational bias of the cells (the fraction of time spent spinning counterclockwise) changed rapidly to a higher stable level, which persisted for the duration of the ramp. The change in bias increased with ramp rate, i.e., with the time rate of change of chemoreceptor occupancy. This behavior can be accounted for by a model for adaptation involving proportional control, in which the flagellar motors respond to an error signal proportional to the difference between the current occupancy and the occupancy averaged over the recent past. Distributions of clockwise and counterclockwise rotation intervals were found to be exponential. This result cannot be explained by a response regular model in which transitions between rotational states are generated by threshold crossings of a regular subject to statistical fluctuation; this mechanism generates distributions with far too many long events. However, the data can be fit by a model in which transitions between rotational states are governed by first-order rate constants. The error signal acts as a bias regulator, controlling the values of these constants.Publication Coordination of Flagella on Filamentous Cells of Escherichia Coli(American Society for Microbiology, 1983) Ishihara, Akira; Segall, Jeffrey E.; Block, Steven M.; Berg, HowardVideo techniques were used to study the coordination of different flagella on single filamentous cells of Escherichia coli. Filamentous, nonseptate cells were produced by introducing a cell division mutation into a strain that was polyhook but otherwise wild type for chemotaxis. Markers for its flagellar motors (ordinary polyhook cells that had been fixed with glutaraldehyde) were attached with antihook antibodies. The markers were driven alternately clockwise and counterclockwise, at angular velocities comparable to those observed when wild-type cells are tethered to glass. The directions of rotation of different markers on the same cell were not correlated; reversals of the flagellar motors occurred asynchronously. The bias of the motors (the fraction of time spent spinning counterclockwise) changed with time. Variations in bias were correlated, provided that the motors were within a few micrometers of one another. Thus, although the directions of rotation of flagellar motors are not controlled by a common intracellular signal, their biases are. This signal appears to have a limited range.Publication Dynamics of Mechanosensing in the Bacterial Flagellar Motor(Proceedings of the National Academy of Sciences, 2013) Lele, Pushkar Prakash; Hosu, Basarab; Berg, HowardMechanosensing by flagella is thought to trigger bacterial swarmer-cell differentiation, an important step in pathogenesis. How flagellar motors sense mechanical stimuli is not known. To study this problem, we suddenly increased the viscous drag on motors by a large factor, from very low loads experienced by motors driving hooks or hooks with short filament stubs, to high loads, experienced by motors driving tethered cells or 1-μm latex beads. From the initial speed (after the load change), we inferred that motors running at very low loads are driven by one or at most two force-generating units. Following the load change, motors gradually adapted by increasing their speeds in a stepwise manner (over a period of a few minutes). Motors initially spun exclusively counterclockwise, but then increased the fraction of time that they spun clockwise over a time span similar to that observed for adaptation in speed. Single-motor total internal reflection fluorescence imaging of YFP–MotB (part of a stator force-generating unit) confirmed that the response to sudden increments in load occurred by the addition of new force-generating units. We estimate that 6–11 force-generating units drive motors at high loads. Wild-type motors and motors locked in the clockwise or counterclockwise state behaved in a similar manner, as did motors in cells deleted for the motor protein gene fliL or for genes in the chemotaxis signaling pathway. Thus, it appears that stators themselves act as dynamic mechanosensors. They change their structure in response to changes in external load. How such changes might impact cellular functions other than motility remains an interesting question.Publication Single-File Diffusion of Flagellin in Flagellar Filaments(Elsevier BV, 2013) Stern, Alan; Berg, HowardA bacterial flagellar filament is a cylindrical crystal of a protein known as flagellin. Flagellin subunits travel from the cytoplasm through a 2 nm axial pore and polymerize at the filament’s distal end. They are supplied by a pump in the cell membrane powered by a proton-motive force. In a recent experiment, it was observed that growth proceeded at a rate of approximately one subunit every 2 s. Here, we asked whether transport of subunits through the pore at this rate could be effected by single-file diffusion, which we simulated by a random walk on a one-dimensional lattice. Assuming that the subunits are α-helical, the answer is yes, by a comfortable margin.