coupled oscillators and biological synchronizationpdodds/files/papers/others/1993/strogatz... ·...
TRANSCRIPT
by Steven H. Strogatz and Ian Stewart
A subtle mathematical threadconnects clocks, ambling elephants,
brain rhythms and the onset of chaos
weight at the end of a string can takeany of an infinite number of closedpaths through phase space, dependingon the height from which it is released.Biological systems (and clock pendulums), in contrast, tend to have not onlya characteristic period hut also a characteristic amplitude. They trace a particular path through phase space, andif some perturbation jolts them out oftheir accustomed rhythm they soon return to their former path. If someonestartles you, say, by shouting, "Boo!",your heart may start pounding but soonrelaxes to its normal behavior.
Oscillators that have a standard waveform and amplitude to which they return after small pertUIbations are knownas limit-eycle oscillators. They incorporate a dissipative mechanism to damposcillations that grow too large and asource of energy to pump up those thatbecome too smali.
THOUSANDS OF FIREFllES Dash in synchrony in this time exposure of a nocturnal mating display. Each insect bas itsown rhythm, but the sight of its neigh·bors' lights brings that rhythm into harmony with those around it. Such couplings among oscillators are at the heartof a wide variety of natural phenomena.
A:ingle oscillator traces out a simple path in phase space. Whentwo or more oscillators are cou
pled, however, the range of possible behaviors becomes much more complex.The equations governing their behaviortend to become intractable. Each oscillator may be coupled only to a few immediate neighbors-as are the neuromuscular oscillators in the smali intestine-or it could be coupled to ali theoscillators in an enormous community. The situation mathematidans find
are espedaliy conspicuous in livingthings: pacemaker cells in the heart; insulin-secreting cells in the pancreas; andneural networks in the brain and spinalcord that control such rhythmic behaviors as breathing, running and che\ving.indeed, not all the oscillators need beconfined to the same organism: consider crickets that chirp in unison and congregations of synchronously flashingfireflies (see "Synchronous Fireflies," byJohn and 8isabeth Buck; SCIENTIFICAMERICAN, May 19761.
Since about 1960, mathematical biologists have been studying simplifiedmodels of coupled oscillators that retain the essence of their biological prototypes. During the past few years, theyhave made rapid progress, thanks tobreakthroughs in computers and computer graphics, collaborations with experimentalists who are open to theory,ideas borrowed from physics and newdevelopments in mathematics itself.
To understand how coupled oscillators work together, one must first understand how one oscillator works byitself. An oscillator is any system thatexecutes periodic behavior. A swingingpendulum, for example, returns to thesame point in space at regular intervals; furthermore, its velodty also risesand falis with (clockwork) regularity.
Instead of just considering an oscillator's behavior over time, mathematidans are interested in its motion throughphase space. Phase space is an abstract.space whose coordinates describe thestate of the system. The motion of apendulum in phase space, for instance,would be drawn by releasing the pendulum at various heights and then plotting its position and velocity. These trajectories in phase space turn out to beclosed curves, because the pendulum,like any other oscillator, repeats thesame motions over and over again.
A simple pendulum consisting of a
Coupled Oscillatorsand Biological Synchronization
102 SCIENTIFIC AMERICAN December 1993
STEVEN H. STROGATZ and IAN STEW·ART work in the middle ground betweenpure and applied mathematics. studyingsuch subjects as chaos and biological oscillators. Strogatz is associate professorof applied mathematics at the Massachusetts Institute of Technology. He earnedhis doctorate at HaIVard University witha thesis on mathematical models of human sleep-Wake cydes and received hisbachelor's degree from Princeton University. Stewart is professor of mathematicsand director of the InterdisdplinaIy Math·ematical Research Programme at the Uni·versity of Warwick in England. He haspublished more than 60 books, including three mathematical comic books inFrench, and writes the bimontWy "~'Iath
cmatical Recreations" column for ScientifiC American.
I n February 1665 the great Dutchphysidst Christiaan Huygens, inventor of the pendulum clock, was con
fined to his room by a minor illness.One day, with nothing better to do, hestared aimlessly at two clocks he badrecently built, which were hanging sideby side. Suddenly he noticed somethingodd: the two pendulums were swingingin perfect synchrony.
He watched them for hours, yet theynever broke step. Then he tried disturbing them-within half an hour they regained synchrony. Huygens suspectedthat the clocks must somehow be influendng each other, perhaps through tinyair movements or imperceptible vibra·tions in their common support. Sureenough, when he moved them to opposite sides of the room, the clocks gradualiy fell out of step, one losing five seconds a day relative to the other.
Huygens's fortuitous observation initiated an entire subbranch of mathematics: the theory of coupled oscillators. Coupled oscillators can be foundthroughout the natural world, but they
EO I £66l "oqWiJJaa NV:mmw :lIJI.I.N3DS
IO~ SC!ENTlnC AMERICAN December 1993
r
age drops instantly to zero-this pattern minlics the firing of a pacemakercell and its subsequent return to baseline. Then the voltage starts rising again,and the cycle begins anew.
A ilistinctive feature of Peskin's model is its physiologically plausible formof pulse coupling. Each oscillator affectsthe others orily when it fires. It kickstheir vollage up by a fixed amount; ifany cell's voltage exceeds the threshold, it fires immeiliately. With these nilesin place, Peskin stated two provocativeconjectures: first, the system would always eventually become synchronized;second, it would synchronize even ifthe oscillators were not quite identical.
When he tried to prove his conjectures, Peskin ran into technical roadblocks. There were no established mathematical procedures for handling arbitrarily large systems of oscillators. Sohe backed off and focused on the simplest possible case: two identical oscillators. Even here the mathematics wasthorny. He restricted the problem further by allowing orily infinitesimal kicksand infinitesimal leakage through theresistor. Now the problem became manageable-ror this spedal case, he provedhis first conjecture.
Peskin's proof relies on an idea introduced by Henri Poincare, a virtuosoFrench mathematician who lived in theearly 1900s. Poincare's concept is themathematical equivalent of stroboscopic photography, Take two identicalpulse-coupled oscillators, A and B, and
attached. Dutch physicist Christiaan Huygens invented thependulwn clock and was the first to observe this phenomenon, inaugurating the study of coupled oscillators.
to handle mathematically because it introduces discontinuous behavior intoan otherwise continuous model and sostymies most of the standard mathematical techniques.
Recently one of us (StTOgatz), along\I;th Renato E. Mirollo of Boston College, created an idealized mathematicalmodel of fireflies and other pulse-coupled oscillator systems. We proved thatunder certain circumstances, oscillatorsstarted at different times will always become synchronized Isee "8ectronic Fireflies," by Wayne Garver and Frank Moss,"The Amateur Scientist," page 1281.
Our work was inspired by an earlierstudy by Charles S. Peskin of New YorkUniversity. In 1975 Peskin proposed ahighiy schematic model of the heart'snatural pacemaker, a cluster of about10,000 cells called the sinoatrial node.He hoped to answer the question of howthese cells synchronize their indMdualelectrical rh\'!hms to generate a normalheartbeat.
Peskin modeled the pacemaker as alarge number of identical oscillators,each coupled equally strongly to all theothers. Each oscillator is based on anelectrical circuit consisting of a capacitor in parallel with a resistor. A constantinput current causes the voltage acrossthe capacitor to increase steadily. As thevoltage rises, the amount of current passing through the resistor increases, andso the rate of increase slows down.When the voltage reaches a threshold,the capacitor discharges, and the volt-
PENDULUM CLOCKS placed near each other soon becomesynchronized (above) by tiny coupling forces transmittedthrough the air or by vibrations in the wall to which they are
easiest to describe arises when each oscillator affects all the others in the system and the force of the coupling increases with the phase difference bet'veen the oscillators. In this case, theinteraction between two oscillators thatare moving in synchrony is minimal.
Indeed, synchrony is the most familiar mode of organization for coupledoscillators. One of the most spectacular examples of this kind of couplingcan be seen along the tidal rivers of Malaysia, Thailand and ew Guinea, wherethousands of male fireflies gather intrees at night and flash on and off inunison in an atlempt to attract the females that cruise overhead. When themales arrive at dusk, their llickerings areuncoordinated. As the night deepens,pockets of synchrony begin to emergeand grow. Eventually whole trees puJsate in a silent, hypnotic concert thatcontinues for hours.
Curiously, even though the fireflies'display demonstrates coupled oscillation on a grand scale, the details of thisbehavior have long resisted mathematical analysis. Fireflies are a paradigm ofa "pulse coupled" oscillator system: tlleyinteract only when onc sees the suddenflash of another and shifts its rhythmaccordingly. Pulse coupling is commonin biology-consider crickets chirpingor neurons communicating via electrical spikes called action potentials-butthe impulsive character of the couplinghas rarely been included in mathematical models. Pulse coupling is awkward
SCIENTIFIC AMERlCAN December 1993 105
POSITION
PHASE PORTRAIT
VELOCITY
eral description of the onset of oscillation. He started by considering systemsthat have a rest point in phase space (asteady state) and seeing what happenedwhen one approximated their motionnear that point by a simple linear function. Equations describing certain systems behave in a peculiar fashion asthe system is driven away from its restpoint. Instead of either returning slowly to equilibrium or moving rapidly out·ward into instability, they oscillate. Thepoint at \vhich this transition takes placeis termed a bifurcation because the system's behavior splits into two branches-an unstable rest state coexists witha stable oscillation_ Hopf proved thatsystems whose linearized form undergoes this type of bifurcation are Iimitcycle oscillators: they have a preferredwaveform and amplitude. Stewart andGolubitsk'Y showed that Hopf's idea canbe c"'<:tended to systems of coupled identical oscillators, whose states undergobifurcations to produce standard patterns of phase locking.
For example, three identical oscillatorscoupled in a ring can be phase-lockedin four basic patterns. All oscillators canmove synchronously; successive oscillators around the ring can move so thattheir phases differ by one third; two oscillators can move synchronously whilethe third moves in an unrelated manner(except that it oscillates with the sameperiod as the others); and two oscillators may be moving half a phase out ofstep, while the third oscillates t\\oce asrapidly as its neighbors.
The strange half-period oscillationsthat occur in the fourth pattern were a
PERJODIC MOTION can be representedin tenns of a time series or a phase portrait. The phase portrait combines position and velocity, thus showing the entirerange of states that a system can display. Any system that undergoes periodic behavior, no matter how complex,will eventually trace out a closed curvein phase space.
TIME SERIES
:~'POSITION:.~ I II I I II I I J
I I I 1~---{,~ '~'---""r-TIME
I I I II I I II I J
I I I '"I I I VELOCITY I
Synchrony is the most obvious caseor a general effect called phaselocking: many oscillators tracing
out the same pattern but not necessarily in step. When two identical osciJIatorsare coupled, there are exactly two possibilities: synchrony, a phase differenceof zero, and antisynchrony, a phase difference of one half. For example, whena kangaroo hops across the Australianoutback, its powerful hind legs oscillateperiodically, and both hit the ground atthe same instant. When a human nmsafter the kangaroo, meanwhile, his legshit the f:,'Tound alternately. If the nehvorkhas more than t\vo oscillators, the rangeof possibilities increases. In 1985 one ofus (Stewart), in collaboration with Martin Golubitsl,y of the University of Houston, developed a mathematical classification of the patterns of networks ofcoupled oscillators, follOWing earlierwork by James C. Alexander of the University of Maryland and Ciles Auchmuty of the University of Houston.
The classification arises from grouptheory (which deals "oth symmerries ina collection of objects) combined withHopf bifurcation (a generalized description of how oscillators "switch on"). In1942 Eberhard Hopf established a gen-
it is not inevilable. Indeed, coupled osdilators often fail to synchronize. Theexplanation is a phenomenon knownas s)"lnmetry breaking, in which a singlesymmetric state-such as synchronyis replaced by several less symmetricstates that together embody the original symmetry. Coupled oscillators area rich source of symmetry breaking.
chart their evolution by taking a snapshot every time A fires.
What does the series of snapshotslook l.ike? A has just fired, so it alwaysappears at zero voltage. The voltage of·B, in contrast, changes from one snapshot to the next. By solving his circuitequations, Peskin found an explicit butmessy formula for the change in B'svoltage between snapshots. The formula revealed that if the voltage is lessthan a certain critical value, it will decrease until it reaches zero, whereas ifit is larger, it will increase. In either case,B \vill eventually end up synchronizedwith A.
There is one exception: if B's voltage is precisely equal to the critical voltage, then it can be driven neither up nordown and so stays poised at criticality. The oscillators fire repeatedly abouthalf a cycle out of phase from eachother. But this equilibrium is unstable,like a pencil balancing on its point. Theslightest nudge tips the system tQ\vardsynchrony.
Despite Peskin's successful analysisof the two-oscillator case, the case ofan arbitrary number of oscillators eluded proof for about 15 years. In 1989Strogatz learned of Peskin's work in abook on biological oscillators by ArthurT. Winfree of the University of Arizona.To gain intuition about the behavior ofPeskin's model, Strogatz wrote a computer program to simulate it for anynumber of identical oscillators, for anykick size and for any amount of leakage.The results were unambiguous: the sys·tern always ended up firing in unison.
Excited by the computer results, Strogatz discussed the problem with Mitollo.They reviewed Peskin's proof of the twooscillatar case and noticed that it couldbe clarified by using a more abstractmodel for the individual oscillators. Thekey feature of the model turned out tabe the slmving upward curve of voltage(or its equivalent) as it rose toward thefiring threshold. Other characteristicswere unimportant.
Mitollo and Strogatz proved that theirgeneralized system always becomessynchronized, for any number of oscillators and for almost all initial conditions. The proof is based on the notionof "absorption"-a shorthand for theidea that if one oscillator kicks anotherover threshold, they "oil remain synchronized forever. They have identicaldynamics, after all, and are identicallycoupled to all the others. The two wereable to show that a sequence of absorptions eventually locks all the oscillatorstogether.
Although synchrony is the simpleststate for coupled identical oscillators,
1
surprise at first, even to Stewart andGolubitsky, but in fact the pattern occurs in rcaltife. A person using a walking stick moves in just this manner:right leg, stick, left leg, stick, repeat. Thethird osciIJator is, in a sense, driven bythe combined effects of the other two:every time one of them hits a peak, itgives the third a push. Because the firsttwo oscillators are predsely antisynchronallS, the third oscillator peaks twicewhile the others each peak once.
The theory of symmetrical Hopf bifurcation makes it possible to classifythe patterns of phase locking for manydifferent networks of coupled oscillators. Indeed, Stewart, in collaborationwith James j. Collins, a biomedical engineer at Boston University, has beeninvestigating the striking analogies between these patterns of phase lockingand the symmetries of animal gaits,such as the trot, pace and gallop.
Quadruped gaits closely resemble thenatural patterns of four· oscillator systems. When a rabbit bounds, for example, it moves its front legs together,then its back legs. There is a phase difference of zero bet\veen the two frontlegs and of one half between the frontand back legs. The pace of a giraffe issimilar, but the front and rear legs oneach side are the ones that move together. When a horse trots, the lockingoccurs in diagonal fashion. An amblingelephant lifts each foot in turn, ,,1thphase differences of one quarter at eachstage. And young gazelles complete thesymmetry group with the prank, a fourlegged leap in which all legs move insynchrony [see "Mathematical Recreations," by Ian Stewart; SClEN1l1'IC AMERICAN, April 1991).
More recently, Stewart and Collinshave extended their analysis to thehexapod motion of insects. The tripod
SYMMETRY BREAKING governs the waysthat coupled oscillators can behave. Synchrony is the most symmetrical singlestate, but as the strength of the coupling between oscillators changes, otherstates may appear. Two oscillators cancouple in either synchronous or antisynchronous fashion (a, b), correspondingroughly to the bipedal locomotion of akangaroo or a person. Three oscillatorscan couple in four ways: synchrony (e),each one third of a cycle out of phasewith the others (d), two synchronousand one with an unrelated phase (e) orin the peculiar rhythm of two oscillators antisyncbronous and the third running twice as fast (n. This pattern is alsothe gait of a person walking slowly withthe aid of a stick.
/
b TWO OUT OF SYNCHRONY
a TWO IN SYNCHRONY
d THREE ONE THIRD OUT OF PHASE
C THREE IN SYNCHRONY
e TWO IN SYNCHRONY AND ONE WILD
f TWO OUT OF SYNCHRONY AND ONE TWiCE AS FAST
106 SCIENTIIX AMERICAN December 1993
d
close to their limit cycles at all times.This insight allowed him to ignore variations in amplitude and to consideronly their variations in phase. To incorporate differences among the oscillators,Winfree made a model that captured theessence of an oscillator community byassuming that their natural frequenciesare distributed according to a narrowprobability function and that in otherrespects the oscillators are identical. Ina final and crucial simplification, he assumed that each oscillator is influencedonly by the collective rhythm producedby all the others. In the case of fireflies,for example, this would mean that eachfirefly responds to the collective nashof the whole population rather than toany individual firefly.
To visualize Winfree's model, imagine a swarm of dots running around acircle. The dots represent the phases ofthe oscillators, and the circle representstheir common limit cycle. If the oscillators were independent, all the dots wouldeventually disperse over the circle, andthe collective rhythm would decay tozero. Incoherence reigns. A simple rulefor interaction among oscillators can restore coherence, however: if an oscillator is ahead of the group, it slows downa bit; if it is behind, it speeds up.
In some cases, this corrective coupling can overcome the differences innatural frequency; in others (such asthat of Gonyaulax), it cannot. Winfreefound that the system's behavior depends on the width of the frequencydistribution. If the spread of frequenciesis large compared ",th the coupling, thesystem always lapses into incoherence,just as if it were not coupled at all. Asthe spread decreases below a criticalvalue, part of the system spontaneously "freezes" into synchrony.
Synchronization emerges cooperatively. If a few oscillators happen tosynchronize, their combined, coherentsignal rises above the background din,exerting a stronger effect on the others.
ahead, and the slower ones fall behind (h, c). A simple coupling force that speeds up slower oscillators and slows downfaster ones, however, can keep them all in phase (d).
c
SCIENTtFlC AMEIUCAN December 1993 107
cies depends on the strength of thecoupling among them. If their interactions are too weak, the oscillatorswill be unable to achieve synchrony. Theresult is incoherence, a cacophony ofoscillations. Even if started in unisonthe oscillators will gradually drift outof phase, as did Huygens's pendulumclocks when placed at opposite ends ofthe room.
Colonies of the bioluminescent algaeGonyaulax demonstrate just this kindof desynchronization. J. Woodland Hastings and his colleagues at Harvard University have found that if a tank full ofGonyaulax is kept in constant dim lightin a laboratory, it exhibits a circadianglow rhythm with a period close to 23hours. As time goes by, the waveformbroadens, and this rhythm graduallydamps out. It appears that the individualcells continue to oscillate, but they driftout of phase because of differences intheir naturai frequencies. The glow ofthe algae themselves does not maintainsynchrony in the absence of light fromthe sun.
In other oscillator communities thecoupling is strong enough to overcomethe inevitable differences in natural frequency. Polymath Norbert Wiener pointed out in the late 1950s that such oscillator communities are ubiquitous inbiology and indeed in all of nature.Wiener tried to develop a mathematicalmodel of collections of oscillators, buthis approach has not turned out to befn.tltful. The theoretical breakthroughcame in 1966, when Winfree, then agraduate student at Princeton UniverSity, began exploring the behavior oflarge populations of limit-cycle oscillators. He used an inspired combination of computer simulations, mathematical analysis and experiments on anarray of 71 electrically coupled neontube oscillators.
Winfree simplified the problem tremendously by pointing out that if oscillators are weakJy coupled, they remain
ba
gait of a coclcroach is a very stable pattern in a ring of six oscillators. A triangle of legs moves in synchrony: frontand back left and middle right; thenthe other three legs are lifted with aphase difference nf nne half.
Why do gaits resemble the naturalpatterns of coupled nscillatnrs in thisway? The mechanical design of animallimbs is unlikely to be the primary reason. Limbs are nor passive mechanicaloscillators but rather complex systemsof bone and muscle controlled by equally complicated nerve assemblies. Themostllkely source of this concordancebetween nature and mathematics is inthe architecture of the circuits in thenervnus system that control locomotion.Biologists have long hypothesized theexistence of networks of coupled neurons they call central pattern generators, but the hypothesis has always beencontroversial. Nevertheless, neurons often act as oscillators, and so, if centralpattern generators exist, it is reasonable to expect their dynamics to resemble those of an oscillator network.
Moreover, symmetry analysis solvesa significant problem in the central-pattern generator hypothesis. Most animals employ several gaits-horses walk,trot, canter and gallop-and biologistshave often assumed that each gait requires a separate pattern generator.Symmetry breaking, however, impliesthat the same central-pattern generatorcircuit can produce all of an animal'sgaits. Only the strength of the couplingsamong neural oscillators need vary.
so far our analysis has been limited to collections of oscillatorsthat are all strictly identical. That
idealization is convenient mathematically, but it ignores the diversity thatis always present in biology. In any realpopulation, some oscillators will alwaysbe inherently faster or slower. The behavior of communities of oscillatorswhose members have differing frequen-
NONIDENTICAL OSCIllATORS may start out in phase with oneanother (as shown on circle a, in which 360 degrees mark oneoscillation), but they lose coherence as the faster Does move
120
of an alignment of molecuies or electronic spins in space.
The analogy to phase transitionsopened a new chapter in statistical mechanics, the study of systems composedof enormous numbers of interactingsubunits, In 1975 Yoshiki Kuramoto ofKyoto University presented an elegantreformulation of Winfree's model, Kuramoto's model has a simpler mathematical slructure that allows it to be analyzed in great detail. Recently Strogatz,along with Mirella and Paul C. Matthewsof the University of Cambridge, foundan unexpected connection bet\veen Kuramoto's model and Landau damping,a puzzling phenomenon that arises inplasma physics when electrostatic wavespropagate through a highly rarefied medium. The connection emerged when we
24
Wben additional oscillators are pulledinto the synchronized nudeus, they amplify its signal, This positive feedbackleads to an accelerating outbreak of synchrony. Some oscillators nonetheless remain unsynchronized because their frequencies are too far from the value atwhich the others have synchronized forthe coupling to pull them in.
In developing his description, Winfree discovered an unexpected link between biology and pbysics. He saw thatmutual synchronization is strikinglyanalogous to a phase transition suchas the freeZing of water or the spontaneous magnetization of a ferromagnet.The width of the oscillators' frequencydistribution plays the same role as doestemperature, and the alignment of oscillator phases in time is the counterpart
~ 72 00TIME IN CONSTANT CONDITIONS (HOURS)
GONYAUlAX luminescent algae (top) change the intensity of their glow accordingto an internal clock that is affected by light. If they are kept in constant dim ligbt,the timing of the glow becomes less precise because the coupling among individual organisms is insufficient to keep them in sync (bottom)_
A sceptical appraisal ofHawking's bestselling
A Brief History of Timerevealing a major fallacy
A SCepticol OPPl"O;sal of hisbestseUing 'A Srief Historyof Time' • revealing Q
major stil'nlifi( folrocy.
'There is a need for freshideas and this work consists
of precisely that.'PROFESSOR JAN BOEYENS
~
/;1H4sLGH4WKII'fGERRED?
GERHARD KRAUS
THE PROFESSIONALHOME WEATHER STATION
Why wail (or a professional weather reportwhen you can have it at your fingertipsanytime you want. The Weather Monitor IIoffers the most complete state-of-the-artweather monitoring system you can buy.
fEATURES INCLUDE: o Highs & lows-Inside & Outside Temps -Instant Melric-Wind Speed & Direction Conversions-Inside Humidity -Outside Humidity•Time & Date & Dew Point Oplion-Barometer '6Rainfall OptionoW'ooo Ch',11 .: ';;¢ -Optional PC
-.. - Interface-Alarms -
WEATHER MONITOR II
Beyourownweathennan.
Inl,oduCliOn byProfcuo, Jon Bocycn.
JANUS PUBLISHING COMPANY£14.99 168pp Hardback Out Now
Available at all good bookshops
Orderloday:l~78-J669 0 SC636BM- F7Ul, 10 S:JO p..m. P.aiic fmt. fAX '-Sl0.6J0.0Sa9
M-t:¥dVlSA· Ont-~ar",'JmInty. JO.d.yIlU'q'-bd;~
Oms l!SImms J46S DoMtoA"., ",,,"AAo,CA94"S
/1. -
aUDla·,:arwm"Room 3005, 96 Broad St.
r.,,;I .... r,J rTn~A'l'7 (')0" A"'L070d
Want tobrush upona foreignlanguage?
With Audio·Forum'sintennediate andadvanced materials, it'seasy to maintain andsharpen your foreign-language skills.
Besides intenncdiate and advancedaudio·cassctte courses-most developedfor the U.S. State Department-we ofTerforeign-language mystery dramas,dialogs recorded in Paris, games, music,and many other helpful materials. And ifyou want to learn a new language, wehave beginning courses for adults andfor children.
We offer introductory and advancedmaterials in most of the world's languages:French, German, Spanish, Italian,Japanese, Mandarin, Greek, Russian,Arabic, Korean, and others. 238 coursesin 86 languages. Our 22nd year.
For FREE 56-page catalog call1-800-662-5180 or write:
Since 1949 more than 15,000authors have chosen the vantagePress subsidy publishing program.You are invited to send for a free illustratedguidebook which explains how your book canbe produced and promoted. Whether your sub
ject is fiction, non·fiction or poetry,scientific, scholarIy, specialized(even controversial), this handsome 32-pagebrochure will showyou how to ar·range for promptsubsidy publica·tion. Unpublished
authors will find this booklet valuable and informative. Foryourfree copy, write to:
VANTAGE PRESS,lnc. Dept. F·53516 W. 34th St., New York, N.Y. 10001
PUBLISHYOUR BOOK
RJRTIlER READING
THE TIMING OF BIOLOGICAL CLOCKS.Arthur T. Winfree. SCientific Americanlibrary, 1987.
fROM Q.OCKS TO CHAOS: 1H:E RHYI1-1MSor liFE. Leon Glass and Michael C.Mackey. Princeton University Press,1988.
SYNCHRONIZATION OF PuLsE-COUPL£DBIOLOGICAL OSCILLATORS. Renata E.MiraDa and Steven H. Strogatz in SlAMJournal on Applied Machematics, Val.50, No.6, pages 1645-1662; December t990.
COuPL£D ONUNEAR OSCIllATORS BELOW TIlE SYNCHRONlZATION 1)-1RESHOW: RElAxATION BY GENERAUZEDlANDAU DAMPING. Steven H. Strogatz,Renata E. MirolJa and Paul C. Matthewsin Physical Review Leners, Vol. 68, No.18, pages 2730-2733; May 4, 1992.
COUPlED NONUNF.AR OSClLLATORS AND
TIlE SYMMETR.IES OF ANIMAL GAITS.J. J. Collins and Ian Stewart in Journalof Nonlinear Science, Vol 3, No.3,pages 349-392; July 1993.
studied the decay to incoherence in oscillator communities in which the freQuency distribution is too broad tosupport synchrony. The loss of coherence, it turns out, is governed by thesame mathematical mechanism as thatcontrolling the decay of waves in such"collisionless" plasmas.
T he theory of coupled osdllatorshas come a long way since Huygens noticed the spontaneous
synchronization of pendulum clocks.Synchronization, apparently a very natural kind of behavior, turns out to beboth surprising and interesting. It isa problem to understand, which is notan obvious consequence of symmetry.Mathematidans have turned to the theory of symmetry breaking to classifythe general patterns that arise whenidentical, ostensibly symmetric oscillators are coupled. Thus, a mathematicaldisdpline that has its most visible rootsin particle physics appears to governthe leap of a gazelle and the ambling ofan elephant. Meanwhile techniques borrowed from statistical mechanics illuminate the behavior of entire populationsof oscillators. It seems amazing thatthere should be a link between the violent world of plasmas, where atoms routinely have thelr electrons stripped off,and the peaceful world of biological osdilators, where fireflies pulse silently along a riverbank. Yet there is a coherent mathematical thread that leadsfrom the simple pendulum to spatialpatterns, waves, chaos and phase transitions. Such is the power of mathematics to reveal the hidden unity of nature.
AT YOURNEWSSTAND
DECEMBER 28
COMINGIN THE
JANUARYISSUE...
Wetland Ecosystems in thelandscape: Fluctuating
Water levels, Biodiversityand Notional Policy
Cyanobacteria and TheirSecondary Metabolites
Breaking Intractability
Animal Sexuality
World languages andHuman Dispersal
TRENDS: CANCEREPIDEMIOLOGY
Timothy M. Beardsley
SEARCHING FORSTRANGE MAnER
Henry J. CrawfordLawrence Berkeley
LaboratoryCarsten Greiner
Duke University
THE FIRST DATANETWORKS
Gerard J. HolzmannAT&TBell Laboratories
Bjorn PehrsonThe Royo/lnstitute of
Technology, Stockholm
ALSO INJANUARY...
SCIENTIFICAMERICAN