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How Microorganisms Move Through Water: the Hydrodynamics of Ciliary Sigma Xi, The Scientific Research Society How Microorganisms Move through Water: The hydrodynamics of ciliary and flagellar propulsion reveal how microorganisms overcome the extreme effect of the viscosity of water Author(s): George T. Yates Reviewed work(s): Source: American Scientist, Vol. 74, No. 4 (July-August 1986), pp. 358-365 Published by: Sigma Xi, The Scientific Research Society Stable URL: http://www.jstor.org/stable/27854249 . Accessed: 18/07/2012 14:21 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Sigma Xi, The Scientific Research Society is collaborating with JSTOR to digitize, preserve and extend access to American Scientist. http://www.jstor.org George T. Yates How Microorganisms Move through Water The hydrodynamicsof ciliaryand flagellar propulsion reveal how microorganisms overcome theextreme effect of the viscosityof water the various The various organisms that propel of and the forces experienced by the fields of engineering, an and themselves through aqueous smallest organisms. physics, applied mathematics, a and environment span over 19 orders of A single drop of water from biology (see for example Gray mass. one or Holwill Wu et al. magnitude in body At pond stream will usually contain Hancock 1955; 1966; extreme are blue whales, the largest hundreds of organisms invisible to 1975; Brennen and Winet 1977). animals ever to inhabit the earth, and the naked eye, a large percentage of are un at the other are microscopic bacteria. which capable of swimming The effects of viscous In terms of absolute the der their own There are more speed, larger power. flows organisms swim faster;when speeds than 50,000 known species of Proto zoa more relative to body size are compared, alone, with new ones being de The larger and familiar free an more as however, microorganisms are the scribed at average rate of swimming animals, such fish and a true champions. Single-celled organ than one day, and most swim at cetaceans, derive their propulsive some water isms, typically between 2 and 1,000 stage of their life cycle (Jahn et thrust by accelerating back (xm long, can sustain swimming al. 1979). Besides being found swim ward, taking advantage of the equal com to speeds up to about 100 body lengths ming freely, microorganisms and opposite inertial reaction force overcome per second. Large pelagic fish like monly occur inside larger organisms, the viscous fluid force or tuna can maintain speeds of only where they exist in symbiotic para which would otherwise slowly arrest are re about 10 body lengths per second, sitic relationships, and some theirmotion. The dominance of iner cases while the world record for human sponsible for disease. tial forces in these is evident maneuvers a swimmers is just over one body Detailed observations of the from the of large ship a an length per second. movements of these tiny organisms approaching harbor. As experi an accom In all cases, organism became possible only after the inven enced mariner will explain, the snip's must or even plishes self-propulsion by putting its tion of the lightmicroscope. The elec power be turned off, or body appendages through period tron microscope has added a new reversed, well before the ship icmovements. After one cycle of the dimension to the study of theirmotil reaches the dock, because the action on motion, the organism and its appen ity, by providing sharp, highly mag of viscosity the hull only gradually mo dages return to their original configu nified fixed images for close examina affects the ship's overwhelming ration, but themean body position is tion of themicrostructures associated mentum. moved forward. The types of appen with locomotion. With the aid of The ratio of inertial to viscous em a dages used, the types of motion these powerful tools, cilia and flagel forces in fluid flows is indicated by ployed, and the very nature of the la have been identified as the cellular nondimensional quantity called the are a fluid forces encountered differ great appendages whose movements Reynolds number. For body of di ly for the various swimmers through most frequently responsible for self mension L moving with velocity U in out a the vast size range. This article propulsion of microorganisms (Fig. viscous fluid, the inertial force is will concentrate on the movements proportional to p(UL)2, the viscous To appreciate fully how propul force varies like (xUL, and the Reyn sion is achieved and olds number is defined as Re = George T. Yates is Senior Scientist in Engineering by ciliary flagel pUL/ lar it is essential to under where is the and the Science at Caltech. He received his B.S. from activity, jx, p density jjl Purdue and hisM.S. and Ph.D. stand some fundamental of University from principles viscosity the surrounding fluid. Caltech. His research on a re or focuses fundamental fluid of hydrodynamics. After brief For fish humans, the Reynolds mechanics, with on we special emphasis view of the relevant principles, number is typically about one mil current biofluiddynamics. His interests include the will look at the structures of cilia and lion, and inertia dominates the fluid mechanics and physiology of swimming and With of both flu motion. As the dimension di extraction winds and flagella. knowledge body flying, optimal energy from id and in hand, we minishes to the scale of currents, and The work dynamics biology microscopic free surface flows. will then be able to discuss the me viscous forces dominate described in this paper has been carried out with organisms, chanics of in detail. This inertial forces, and the support from theNational Science Foundation and propulsion Reynolds a subdivision of number becomes 0.1 or smaller. At theOffice ofNaval Research. Address: subject, biofluiddyn is its nature interdisci the even smaller dimension of the Engineering Science, Caltech, Pasadena, CA amics, by very 91125. plinary; it requires collaboration by individual cilia and flagella, which 358 American Scientist, Volume 74 oscillate typically between 10 and 30 cycles per second, the Reynolds number is usually less than 10~5. Because the inertial effects are of no significance at these small scales of motion, microorganisms are unable to take advantage of the inertial reac tion and must derive their forward thrust from viscous reactions. The typical viscous forces which act on a human swimmer are at least ' 6 orders of magnitude smaller than _ i the inertial forces. To the 1 ji.m appreciate nature of the viscous forces experi enced by microorganisms, we would have to swim in a fluid one million times more viscous than water. Even a in pool filled with honey the rela tive strength of viscous to inertial forces would be two orders ofmagni tude smaller than it is formicroorga nisms swimming in water, and a more appropriate fluid would be mo lasses or even tar. Whenever the Reynolds number is small, the rate of change in mo over mentum time can be neglected relative to the viscous and pressure components, and time becomes sim ply a parameter. This means that the fluid motion generated in response a to microorganism's movement de pends on its instantaneous velocity, and not on its velocity at any previ ous instant or on the rate of change of any other quantity with time. Fur thermore, because the fluid inertia (mass times acceleration) is negligibly small, any external force must be balanced by the viscous stress and the pressure in the fluid. can now We begin to appreciate the difficulties encountered in achiev ing self-propulsion at low Reynolds numbers. For a swimmer?for exam 1. Cilia and are the Figure flagella ple, a human in an ocean of molas appendages that propel microorganisms ses?who moves his arms from a water. are from through They distinguished to a final con each other by their length relative to the cell given starting position as the instantaneous fluid body, well as by their structures and figuration, are characteristic movements. Flagella often force depends linearly on the velocity are attached to one end of the body, and of the motions. When the more than twice the of the body frequently length swimmer not body. The bacterium Salmonella abortus stops moving, only a does his absolute motion but all equi (top) has many flagella, which form stop, bundle that is behind the 10 comes clearly visible jJLlTl the surrounding fluid to rest cell in this electron The bundle and thus body micrograph. intermittently unwraps reforms, immediately, because there are no a sea Ciona is changing the swimming direction. A spermatozoon from squirt (bottom left) inertial effects. Furthermore, if the to 200 flashes shown swimming in sea water in this series of flash photographs (top bottom, swimmer retraces the taken with a The wave is and to exactly paths per second) light microscope. propulsive planar propagates followed his arms to their initial the the the cell moves to the left. Cilia (bottom by right along flagellum. Consequently, right), he will find himself are numerous in rows on the configuration, which are usually much shorter than the body, and arranged the the cell (and the in the same body surface. Gradual variation in the phase of the beat cycle along length of fluid) exactly fromwhich he started.
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