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FOCUS ISSUE: The constructive role of noise in fluctuation driven transport and stochastic resonance Overview: The constructive role of noise in fluctuation driven transport and stochastic resonance
R. Dean Astumian
Departments of Surgery and of Biochemistry and Molecular Biology, MC 6035, University of Chicago, Chicago, Illinois 60637

Frank Moss
Center for Neurodynamics, University of Missouri at St. Louis, St. Louis, Missouri 63121

Received 17 July 1998; accepted for publication 17 July 1998 Random noise is typically thought of as the enemy of order rather than as a constructive influence. Recent work has shown however that under certain circumstances, noise and Brownian motion can facilitate transmission of information via a mechanism know as stochastic resonance, and help systems use chemical energy and nonequilibrium fluctuations to drive directed motion via fluctuation driven transport. In this focus issue we have collected several articles that capture the flavor of these developing fields and point the way to new directions for research. © 1998 American Institute of Physics. S1054-1500 98 02203-4

Since the development of a statistical description of thermodynamics by Maxwell, Boltzmann, and many others, scientists have attempted to devise ways to harness the fluctuations inherent even in equilibrium systems to accomplish work. While no perpetual motion machines are on the market nor are any expected to appear the thought experiments involved have provided excellent theoretical laboratories in which to investigate the deeper implications of the second law of thermodynamics. Perhaps the first and most influential such construct is that of Maxwell, who proposed a device often whimsically described as an intelligent being or demon that could open or close a gate separating two containers of a gas depending on a measurement of the velocity of an oncoming molecule.1,2 If the demon would open the gate only to fast molecules coming from one side and to slow molecules coming from the other, a thermal gradient would be formed. Subsequent analysis has focused on understanding the energy required to make, store, and eventually erase the necessary measurements of the molecular velocities. Energy dissipation can arise in any of these processes and the minimum amount of energy dissipated is equal to or greater than the work stored by the creation of a thermal gradient.3 Maxwell’s demon requires active acquisition of information for operation. Macroscopic ratchets and rectifiers, however, passively convert external zero average fluctuating forces into net motion current . What happens if such devices are shrunk to microscopic size? This was the question posed by Smoluchowski,4 and later by Feynman in his Lectures on Physics,5 where he analyzed a microscopic ratchet a cog with asymmetric teeth and pawl Fig. 1 a . The ratchet’s teeth are arranged so that it is impossible to drag the pawl up one face. At first glance it would seem that a paddle1054-1500/98/8(3)/533/6/$15.00 533

wheel attached to such a ratchet should convert thermal fluctuations to unidirectional motion. If the wheel cannot go backward, molecules hitting the paddle would cause an irregular but relentless rotation of the wheel that could lift a weight. This would be a perpetual motion machine of the second kind, a contradiction to the second law. Feynman resolved this paradox, showing that consistent application of the laws of statistical mechanics to all parts of the device restores the result that work cannot be extracted from thermal noise in an isothermal system. In order to function, the pawl must be attached to the ratchet by some elastic element, say a spring, which is also influenced by thermal noise. When the pawl is down, the device works as anticipated. However, occasionally the pawl is up, and is disengaged from the ratchet cog. In this rare event, molecular collisions with the paddlewheel cause forward and backward motion with equal probability. Because of the asymmetry, it takes only a tiny motion backward to set the device back by one tooth, but a much larger movement forward to advance the device by one tooth. If the paddlewheel and pawl are at the same temperature, the tendencies to move forward, due to the fluctuating force acting at the paddlewheel, and to move backward due to the fluctuating engagement and disengagement of the pawl, exactly cancel, and despite appearances, the ratchet will not rotate. Recently, Kelly and colleagues at Boston College6–8 have synthesized an organo-molecular ratchet with a triptycene as the paddlewheel, linked to a four ring helicene as the pawl and spring Fig. 2 . Because the helicene forms a helix, the force necessary to turn the triptycene paddlewheel clockwise is less than is necessary to turn it counterclockwise. This is illustrated by manipulation of ‘‘tinker-toy’’ molecular models and by the calculated plot of H versus the dihedral angle. The energy profile is strongly anisotropic as
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Astonishingly. Astumian and F. consistent with the second law. Feynman showed that when the temperatures are different the imbalance in the strength of the thermal noise acting on the paddlewheel and on the spring does indeed cause the cog to rotate. U 50 meV. the differential equation describing the time evolution of the system is dn dt dn dt W W t n t W W t t n W t n . a Depiction of Feynman’s ratchet. However. 1998 R. simple equation illustrating this can be derived from the two relations for the unidirectional rates shown in Fig. When the spring is down. 1. the random molecular forces on the paddle cause forward and backward motion with equal probability.29 see also the papers in this volume by Wiesenfeld and Jaramillo. Binding substrate at the active site reduces the flexibility of the pawl. The net rate is nonmonotonic with respect to the average temperature T Fig. with T 1 T 2 2 T and T 1 T 2 2T. Despite the anisotropy in the structural design. at equilibrium. with A 104/s. the ratchet’s rotation is counterclockwise! A where k B is Boltzmann’s constant. When the active site is empty. Vol. Certainly in the case of the molecular ratchet in Fig. it was shown that despite the anisotropy. If the catalyzed reaction is far from equilibrium the fluctuations of the barrier height will not obey detailed balance and can lead to unidirectional rotation of the triptycyl paddlewheel—a chemically driven molecular motor. The equations are approximate formulas for the frequency of moving a step in the clockwise cw and counterclockwise ccw direction. but of course is periodic with a repeat every 120°. or to a moiety that absorbs light and thus modulates the height of the barrier for rotation? Recent research. Feynman also considered a modification of this ratchet. 3. whereas to send it forward by a tooth requires a much greater motion. 3 a 13. and raises the barrier to rotation. Using the NMR technique of spin polarization transfer. 2. illustrating the synergy between ‘‘thermal noise’’ and an external energy supply the thermal gradient in this case for doing work. It only takes a very tiny movement of the wheel backwards to set the device back one tooth. for structural reasons analogous to those given by Feynman for his ratchet and pawl. despite appearances will not turn.534 Chaos. 2 Downloaded 22 Feb 2006 to 130.28. For generality we have described a system with two pathways between the states. such as linking the helicene pawl to a chemical catalyst. the ratchet rotates clockwise as our intuition would suggest. Redistribution subject to AIP license or copyright. Moss FIG. disengaging the oneway mechanism. h 2 where is the spring constant and h is the height of a tooth on the U cog. Because of the asymmetry of the ratchet’s teeth. and T T: net rate A U T e 2k B T 2 U/k B T . 1. c Hypothetical scheme for converting this ratchet to a chemically driven molecular motor by linking the ‘‘pawl’’ to a catalyzed reaction conversion of S to P . in the contexts of stochastic resonance SR 9–26 and fluctuation driven transport. D. the device works both ways—when the temperature of the paddlewheel is greater than that of the spring. 1 expected for a ratchet. The temperatures of the pawl and of the gas in the box containing the paddlewheel need not be equal. 1 b . and the barrier to rotation small. FIG. If the paddle and the pawl are at the same temperature so that the fluctuations on the pawl are as strong as those driving the paddlewheel.163. it might seem that even thermal fluctuations acting on a paddlewheel attached to the ratchet could be used to do work. but when the temperature of the spring is greater than that of the paddlewheel. 8. EARLY THEORETICAL MODELS Early theoretical work for both SR and fluctuation driven transport focused on a two-state model such as shown in Fig. However. the frequencies of clockwise and counterclockwise turns are exactly equal—a neat experimental verification of Feynman’s theoretical analysis. and by Fulinski . molecular collisions with the paddlewheel indeed tend to cause the cog to turn only in the planned direction. see http://chaos. clockwise and counterclockwise transitions over the barrier are equally likely. No. both of which can serve as an energy source. in the rare event that the spring is the behavior of the spring is also influenced by collisions with molecules which cause it to vibrate. b Plot of net rate vs T from Eq. Is there some other possibility. b Calculated H as a function of the dihedral angle about this bond. If n i is the probability for the system to be in state i. and T 10 K. and is maximized when U 2k B T. the ratchet. the pawl is flexible.jsp . 2 there is no way to power it with a thermal gradient. II. a A molecular ratchet constructed of a triptycyl paddlewheel attached to a helix forming helicene pawl by a single bond around which rotation is possible.108.27–42 has focused on the interplay between thermal noise and catalyzed chemical reactions or external signals.aip. 1 .

McNamara and Wiesenfeld13 focused on the correlation between the applied field and occupancy of the ‘‘ ’’ and ‘‘ ’’ states. see http://chaos. D. 3 a is identical to the Michaelis–Menten mechanism for an enzyme catalyzed reaction31 E S ES E P.111. chemical kinetic model. It is worth noting that the two state model in Fig. x 0 is an offset to position the maxima at 0. the symmetric case with only one pathway between the states need be considered. Cyclic flux occurs despite the fact that the chemical driving force k B T ln( 1 2 / 1 2) 0 at every instant in time notwithstanding the perturbation of the energy levels. If the chemical potential of P is greater than that of S. Moss 535 i t 0 i exp 1 i U t /D for i 1. D. but for U(t) U 0 cos( t) the stationary oscillating solution can be expanded as a Fourier series and an explicit recursion relation given for the coefficients for all harmonics. . Vol.28 This is easily extended to systems with an arbitrary number of states. Eq. b Illustration of how fluctuation or oscillation of the relative energies of the two states can give rise to cyclic flux. Under most circumstances. Astumian and F. 1998 R.2. where U(x) is plotted directly vs x. Kinetically. 5 . No. 3. where D is the amplitude of white noise. 5 where the free ‘‘E’’ is the ‘‘ ’’ state.163.29 analyzed the net flux around the cycle caused by the oscillation of the energy levels of the two states.2 to vary with time i t 0 i exp i U t /D . Astumian and colleagues28. The best system performance is achieved with an optimum noise intensity. When the noise intensities are too small they cause the signal to be barely or Downloaded 22 Feb 2006 to with U(x) U 0 (x) cos2(x/2)] the ball most likely flows from the ‘‘ ’’ to ‘‘ ’’ state also via the clockwise path. the amplitude of this flux is a nonmonotonic function of the noise. The case with 1 1 and 2 0 however. 3. etc.jsp . 2 . The thermodynamic efficiency can be appreciable. thus undergoing a net clockwise cycle. the barriers for paths 1 and 2 are each equally spaced between the ‘‘ ’’ and ‘‘ ’’ states. Redistribution subject to AIP license or copyright. Focusing on fluctuation driven transport. describes a situation with a gentle slope always in the clockwise direction see Fig. When the energy of the ‘‘ ’’ state is low left-hand plot.sin( x). and the concentrations of S and P are assumed to vary slowly. and using an oscillating or fluctuating modulation of the energy levels of the two states to drive clockwise flow means that energy is transduced from the non-equilibrium fluctuation to do work on the chemical reaction S P. and where 3 t t 2 2 t . and bound ‘‘ES’’ is the ‘‘ ’’ state. but can in principle be externally controlled.108. The 8 2 potential is U 0 (x) i 1 exp( i /9)sin„ 2i(x x 0 )…/i which is modu2 lated by adding cos (x/2). thus opening the field to a much wider range of experimental tests.’’ to a weak information carrying signal can enhance the signals detectability by some nonlinear system or intensify the information content of the system’s output.28 Recent work has gone far toward relaxing the restrictions implicit in this adiabatic. FIG. III.43 In the case of stochastic resonance. When the energy of the ‘‘ ’’ state is low right-hand plot. with U(x) U 0 (x) cos2(x/2)] the ball most likely flows from ‘‘ ’’ to ‘‘ ’’ in a clockwise direction. We show parametric plots of cos( x). c An alternative view. this is identical to the mechanism for a Michaelis– Menten enzyme. In this picture. They showed that the signalto-noise ratio calculated from the power spectrum depends nonmonotonically on the value of the white noise amplitude. 8. Equations 3 can be formally solved in terms of integrals for any time dependence of the rate coefficients. the mechanism for generating left to right motion is apparent. a A two state kinetic mechanism with two pathways between the states. D k B T according to the fluctuation dissipation theorem.U(x) .aip. 4 Stochastic resonance SR is a term which has been widely adopted to describe processes whereby the addition of a random function. In the symmetric case. 1 2 1/2. often called the ‘‘noise. For this purpose. As with SR. 3 . The apportionment constant i reflects the symmetry of the barrier relative to the two states. there is a counterclockwise torque on the cycle.Chaos. t . STOCHASTIC RESONANCE using the conservation relation n W W t t 1 1 n 1. A time-dependent signal that perturbs the relative energies of the ‘‘ ’’ and ‘‘ ’’ states such that (U (t) U (t)) U(t) will also cause the transition coefficients 1.

energy barriers of roughly 10k B T hydrolysis of one molecule of ATP releases about 20k B T and a viscosity Downloaded 22 Feb 2006 to 130. it has been shown that not even a hard threshold is necessary. The concept was first introduced nearly 20 years ago. and depends on the statistical properties of the fluctuation as well as on the anisotropy of the potential. These ideas can also be applied to quantum systems51 see the paper by Riemann and Hanggi in this issue .36 used an optical trap to show the same behavior at the level of a single particle. Huxley44 proposed a model for muscle movement based on biased Brownian motion. The direction of the flux induced can reverse. 8. No. and molecular versions of other macromechanical devices. Astumian and Bier33 and Prost et al.18 Several reviews19–21 and popular articles22–24 have been written. A very significant recent development was the demonstration by Ajdari and Prost30 that directed motion of a particle in a viscous medium could be induced simply by turning on and off a periodic anisotropic potential.163. One of the main reasons for the tremendous recent interest in fluctuation driven transport has been the potential applications to biological systems. and colleagues46. could also show the effect. These results triggered an avalanche of research that persists to this day. with two specific topics coming to the fore—applications to particle separation and possible ties to the mechanism of biomolecular motors and pumps.41 dealing with fluctuation driven transport. They have argued that this can also be used as the basis for a continuous separation technique. Breymeyer. When they are too large. D. Several authors55–57 have shown that with appropriate potentials there can be a threshold value of the diffusion coefficient where particles with a diffusion coefficient greater than the threshold move in a direction opposite to those with a diffusion coefficient smaller than the threshold.34 showed that this mechanism now known as a flashing ratchet works even if the potential or equivalently the interaction is turned on and off randomly.61 Molecular motors use chemical energy to move along a biopolymeric track constructed of identical monomeric subunits. a notable turning point came when it was recognized that simple threshold systems.54 Rousellet35 demonstrated experimentally that diffusion of small particles can be biased by turning an anisotropic dielectrophoretic potential cyclically on and off.536 Chaos. Vol. Analysis of the simplest flashing ratchet models shows that with a spatial period of 10 8 m. Risken45 showed that the mobility of a particle in an anisotropic periodic potential is not symmetric with respect to an applied force. with a velocity and sign that depends on the phase relation between the temperature and potential periods. brakes.53. If the system has a nondiagonal mobility Shortly after. Moss not at all detectable.108. described only by statistics. This mechanism is termed a ‘‘rocking ratchet’’ 49 because the force can be visualized as randomly rocking the potential back and forth. Magnasco32 showed that a zero average slowly fluctuating macroscopic force causes directed motion of a small particle moving along an anisotropic potential. and inorganic phosphate Pi to depend on the position of a motor molecule myosin along a biopolymer actin . This opens up the possibility of a continuous separation. adenosine diphosphate ADP . IV.aip. 3. Perhaps a third topic should now be added to this list—chemically synthesized nanomolecular ratchets. 1998 R.47 investigated the effect of harmonic forcing on particles in a spatially periodic pinning potential. Passive devices that can convert external random fluctuations into directed motion have also been investigated. Originally proposed as a possible method for particle separation this represents a true Brownian motor because the mechanism utterly fails in the abscence of thermal noise.26 These generalizations on the conditions which systems must satisfy in order to show SR have stimulated new applications in a much wider class of systems. see http://chaos. Doering et al.60 particularly in light of the fact that it is now possible to study biomolecular motors at the single molecule level. in the context of global meteorology. FLUCTUATION DRIVEN TRANSPORT The phenomenon of fluctuation driven transport was anticipated in some sense by many authors.52 There have been several recent reviews37–39 and popular articles40.111. Even before publication of Feynman’s ratchet.jsp .10 It was demonstrated in physical systems for the first time using an electronic circuit analog.25 More recently. Because of this the motors see an energy landscape that is periodic at every instant in time. Wonneberger. Though SR was originally thought to exist only in dynamical systems described by bi.15. where particles are continuously fed into the middle of the device and large particles collect on one side and small particles collect on the other side. and the flashing ratchet models are thus more relevant than the fluctuating force ratchets. but only within a frequency window for the inverse correlation time of the fluctuation.50 analyzed the transition to the very high frequency regime and demonstrated that the behavior in this region is very rich. irrespective of the amplitude of the potential. Astumian and F. This work has begun an avalanche of proposals concerning ways to use fluctuation driven transport for particle separation. as a theory of the recurrences of the Earth’s Ice Ages. Buttiker48 showed that a symmetric spatially periodic temperature could drive transport of a Brownian particle along a spatially periodic potential. Faucheaux et al. Recent work has dramatically extended the domain of fluctuation driven transport. some having invaded such far-flung fields as biology14 and medicine. randomness overwhelms the information content.7. His idea in some ways is very similar to Maxwell’s demon because it requires the rate constants for binding and release of adenosine triphosphate ATP .9. and to systems with intertia. a dc force in one direction will cause transport perpendicular to the force. These experimental setups have used a flashing ratchet approach.16.or multistable energy potentials. Redistribution subject to AIP license or copyright. Closer in spirit to Feynman’s work. Duke and Austin58 and Ertas59 have recently proposed a novel two-dimensional separation method based on constructing a system with anisotropic barriers using microlithography.11 The subject then lay dormant until the late ’80s when a remarkable experiment was performed with a bistable ring laser12 demonstrating the phenomenon of stochastic resonance with an experimentally controllable system.

19 F. Buckley.62. D. 770 1996 . 909 1998 . 5 1983 . P. S. von Baeyer. P. D. Chiou-Tan.65 or randomly fluctuating66. Engl. who discuss SR in mammalian brain tissue. . J. Reading. the velocity induced by turning the potential on and off 200 times per second is around 10 6 m/s and a force of about 1–10 pN is necessary to bring the particle to a halt. Chem. edited by G. I.67 electric field to drive unidirectional transport. Astumian and F. Feynman. and J. Relying on collaboration between physicists. ‘‘Theory of stochastic resonance. 205. O’Gorman.’’ Nature London 393. 8 A. and H. the barrier to rotation is low and the brake is off. Heslot. Sestelo. R. Benzi.’’ J. it might also be possible to modulate the barrier height between a photochemically excited state and a ground state. Spatio-temporal SR is demonstrated in glial cell networks and in the Belousov–Zhabotinsky chemical reaction by Peter Jung. ‘‘New molecular devices: in search of molecular ratchets. the last two articles give some exciting new experimental results on motion driven by ratchets Gorre-Tallini et al. 1990 . With an appropriate photoactive moiety. and F. and R. ‘‘Demons. ‘‘Observation of stochastic resonance in a ring laser. Moss. ‘‘In search of molecular ratchets. When a metal ion is bound to the chelator. Redistribution subject to AIP license or copyright. T. C. 769 1996 . McNamara. if the catalyzed reaction would be far from equilibrium. Am. Tellitu. Bifurcation Chaos Appl. T. P. 3 C. Chem. Finally. 16 P. 12 B. Vulpiani. Collins and his collaborators move the subject into the realm Downloaded 22 Feb 2006 to 130. the polycycle is rigid. While this issues coverage of current work on SR and fluctuation driven transport is certainly not complete. and biologists. and F. Hunt and colleagues then present an experimental and theoretical work on spatio-temporal SR using a chain of coupled electronic circuits. T. T. J. L.’’ Angew. Moss. and quantum SR and Brownian motors Reimann and Hanggi . Sci. Cordo. Sands.’’ Angew. Sutera. R.’’ Sci. Davis. noise and chaos mediated encryption Minai and Pandian . energy from the chemical reaction would drive counterclockwise! unidirectional rotation of the triptycyl paddlewheel. Lutchen. 2 H. 5 R. Merfeld. and a number of authors have described more complicated models that take into account more degrees of freedom for the system. with many new approaches and applications emerging.63 Perhaps the strongest direct evidence for a ratchet mechanism for free energy transduction by a biomolecule comes from recent experiments showing that the Na. Moss. Ann Cornell-Bell. H. such simple models do not reflect all aspects of the very complicated motor proteins. we can look toward harnessing noise to help construct microscopic motors and pumps. Moss. These results were interpreted theoretically in terms of a four state kinetic ratchet. Sci. Pierson.’’ Phys. 1998 R. J. Vol. 127 1998 . P.163. Information. and the Second Law.43 V. ‘‘Long term climatic transitions and stochastic resonance. Computing Princeton University Press. ‘‘Stochastic Resonance. Alencar. see http://chaos. 15 F. After an admirable minireview of stochastic resonance by Kurt Wiesenfeld and Fernan Jaramillo. P. F. These values are in good agreement with what is observed experimentally. Rev. J. we hope it is representative of some of the challenging and intriguing research areas wherein noise enhancements can usefully be sought. ‘‘Experimentell nachweisbare der ublichen thermodynamik wiedersprechende molekularphanomene. 3655 1998 .’’ Tellus 34. ‘‘Stochastic resonance in a bistable system. ‘‘Stochastic resonance: Tutorial and update. 8. Sestelo. A 4854 1989 . . Grigg. ‘‘The mechanism of stochastic resonance. Leff and A. A 97. Kelly and colleagues previously synthesized a molecular brake. S. ‘‘Life-support system benefits from noise. A. Then.’’ but when the chelator is not bound to a metal ion. MA. Org. Rex. Pantazelou. 21 L. The fields of SR and fluctuation driven transport are actively developing. Schiff and his co-workers.’’ J. 337 1993 . NY. 18 B. 1069 1912 . P. 60. Chem. S.’’ Phys. 14 J. 20 F. Bennet. J.108. Zapperi. Wiesenfeld. S. Robinson. J. S. A similar strategy could be adopted in the case of the ratchet. H. Jung. Tellitu. S. E. 3. 36. we begin to see a way that chemical or light energy can be used to power a simple molecular motor. Mod. Hanggi. 1994 . B. and J.aip. 223 1998 . 1998 . In this focus issue we provide a sampling of some of these new applications. 6.’’ Phys. E. Finally. Wiesenfeld.111.’’ Nature London 383. Phys.Chaos. but with a catalyst in place of the simple chelator. ‘‘Tilting at windmills? The second law survives.’’ Nature London 365. I.’’ Int. and P. M. Douglass. Inglis. Eng. Z. Continuing the biological network theme is S. Kelly. and Derenyi et al. P. 1 Returning to consideration of the molecular ratchet in Fig. Suki. K. Int. a biomolecular ion pump can use an external oscillating64. and F. and Ken Showalter and his colleagues. Rev. we are led through topics in SR with aperiodic stimuli Collins et al.’’ Phys. Coupled with the intrinsic anisotropy of the system. Collins. Needless to say. Tuel. Engines. Bifurcation Chaos Appl.’’ in Contemporary Problems in Statistical Physics. Int. ‘‘Enhancement of subthreshold sensory nerve action potentials during muscle tension mediated noise. K. Magee. No. Stanley. Y. New York. Ed. J. J. 1 1982 . NJ.jsp . ‘‘Noise enhancement of information transfer in crayfish mechanoreceptors by stochastic resonance. 13. Leighton. Fulinski discusses fluctuating barriers and membranes. Lett. 10 C. K. Gammaitoni. and D. and F. Phys. presenting a large barrier to rotation of the triptycyl paddlewheel—the brake is ‘‘on. Nelson. ‘‘Noise-enhanced tactile sensation. M. 6 T. Moss 537 somewhat greater than that of water. The Feynman Lectures on Physics Addison-Wesley. Moss. 11 S. Collins. S. Cell membranes are again the topic when SR at the molecular level is discussed by Sergey Bezrukov and Igor Vodyanoy. chemical energy would drive a nonequilibrium fluctuation between a high barrier and low barrier—a flashing ratchet. 1383 1994 . 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Collins. E. The richness of the high frequency behavior of a rocked ratchet is explored mathematically by Doering et al. Verschueren. PERSPECTIVE of psychophysics with a beautiful experiment on the human tactile system.

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