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Life at Low Reynolds Number

Life at Low Reynolds Number

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05/13/2014

 
LifeatlowReynoldsnumber
E.M.Purcell
LymanLaboratory,HarvardUniversity,Cambridge,Massachusetts02138
(Received12June1976)
Editor'snote:
Thisisareprint(slightlyedited)ofapaperofthesametitlethatappearedinthebook
PhysicsandOurWorld:ASymposiuminHonorofVictor
F.
Weisskopf,
publishedbytheAmericanInstituteofPhysics(1976).Thepersonaltoneoftheoriginaltalkhasbeenpreservedinthepaper,whichwasitselfaslightlyeditedtranscriptofatape.Thefiguresreproducetransparenciesusedinthetalk.Thedemonstrationinvolvedatallrectangulartransparentvesselofcornsyrup,projectedbyanoverheadprojectorturnedonitsside.Someessentialhandwavingcouldnotbereproduced.ThisisatalkthatIwouldnot,I'mafraid,havethenervetogiveunderanyothercircumstances.It'sastoryI'vebeensavinguptotellViki.Likesomanyofyouhere,I'veenjoyedfromtimetotimethewonderfulexperienceofexploringwithVikisomepartofphysics,oranythingtowhichwecanapplyphysics.Wewanderaroundstrictlyasamateursequippedonlywithsomeelementaryphysics,andintheend,itturnsout,weimproveourunderstandingoftheelemen-taryphysicsevenifwedon'tthrowmuchlightontheothersubjects.Nowthisisthatkindofasubject,butIhavestillanotherreasonforwantingto,asitwere,needleVikiwithit,becauseI'mgoingtotalkforawhileaboutviscosity.ViscosityinaliquidwillbethedominantthemehereandyouknowViki'sprogramofexplainingeverything,in-cludingtheheightsofmountains,withtheelementaryconstants.Theviscosityofaliquidisaverytoughnuttocrack,ashewellknows,becausewhenthestuffiscooledbymerely40degrees,itsviscositycanchangebyafactorofamillion.IwasreallyamazedbyfluidviscosityintheearlydaysofNMR,whenitturnedoutthatglycerinewasjustwhatweneededtoexplorethebehaviorofspinrelax-ation.Andyetifyouwerealittlebuginsidetheglycerine,lookingaround,youwouldn'tseemuchchangeinyoursurroundingsastheglycerinecooled.Vikiwillsaythathecanatleastpredictthe
logarithm
oftheviscosity.Andthat,ofcourse,iscorrectbecausethereasonviscositychangesisthatit'sgotoneoftheseactivationenergythingsandwhathecanpredictistheorderofmagnitudeoftheexponent.Butit'smoremysteriousthanthat,Viki,becauseifyoulookattheChemicalRubberHandbooktableyouwillfindthatthereisalmostnoliquidwithviscositymuchlowerthanthatofwater.Theviscositieshaveabigrange
buttheystopatthesameplace.
Idon'tunderstandthat.That'swhatI'mleavingforhim.'Now,I'mgoingtotalkaboutaworldwhich,asphysicists,wealmostneverthinkabout.Thephysicisthearsaboutviscosityinhighschoolwhenhe'srepeatingMillikan'soildropexperimentandheneverhearsaboutitagain,atleastnotinwhatIteach.AndReynolds'snumber,ofcourse,issomethingfortheengineers.Andthe
low
Reynoldsnumberregimemostengineersaren'teveninterestedin-exceptpossiblychemicalengineers,inconnectionwithfluidizedbeds,afascinatingtopicIheardaboutfromachemicalengineeringfriendatMIT.ButIwanttotakeyouintotheworldofverylowReynoldsnumber-aworldwhichisin-habitedbytheoverwhelmingmajorityoftheorganismsinthisroom.Thisworldisquitedifferentfromtheonethatwehavedevelopedourintuitionsin.Imightsaywhatgotmeintothis.Tointroducesomething
3
AmericanJournalofPhysics,Vol.45,No.1,January1977
thatwillcomelater,I'mgoingtotalkpartlyabouthowmicroorganismsswim.Thatwillnot,however,turnouttobetheonlyimportantquestionaboutthem.IgotintothisthroughtheworkofaformercolleagueofmineatHarvard,HowardBerg.BerggothisPh.D.withNormanRamsey,workingonahydrogenmaser,andthenhewentbackintobiologywhichhadbeenhisearlylove,andintocellularphysiology.HeisnowattheUniversityofColoradoatBoulder,andhasrecentlyparticipatedinwhatseemstomeoneofthemostastonishingdiscoveriesaboutthequestionswe'regoingtotalkabout.SoitwaspartlyHoward'swork,tracking
E.coli
andfindingoutthisstrangethingaboutthem,thatgotmethinkingaboutthiselementaryphysicsstuff.Well,herewego.InFig.1,youseeanobjectwhichismovingthroughafluidwithvelocity
v.
Ithasdimension
a.
InStokes'slaw,theobjectisasphere,buthereit'sanything;
71
and
p
aretheviscosityanddensityofthefluid.Theratiooftheinertialforcestotheviscousforces,asOsborneReynoldspointedoutslightlylessthanahundredyearsago,isgivenby
avp/7J
or
aviv,
where
u
iscalledthe
kinematic
viscosity.It'seasiertorememberitsdimensions:forwater,
v""
10-
2
cm
2
/sec.TheratioiscalledtheReynoldsnumberandwhenthatnumberissmalltheviscousforcesdominate.Nowthereisaneasyway,whichIdidn'trealizeatfirst,toseewhoshouldbeinterestedinsmallReynoldsnumbers.
If
youtaketheviscosity
71
andsquareitanddividebythedensity,yougetaforce(Fig.2).Nootherdimensionscomeinatall.
71
2/
P
isaforce.Forwater,since
71""
1
0-
2
and
p""
1,71
2/
p""
10-
4
dyn.Thatisaforcethatwilltow
anything,
largeorsmall,withaReynoldsnumberoforderofmagni-tude1.Inotherwords,ifyouwanttotowasubmarinewithReynoldsnumber1(orstrictlyspeaking,1/67rifit'sasphericalsubmarine)towitwith
10-
4
dyn.Soit'sclearinthiscasethatyou'reinterestedinsmallReynoldsnumberifyou'reinterestedin
smallforces
inanabsolutesense.TheonlyotherpeoplewhoareinterestedinlowReynoldsnumber,althoughtheyusuallydon'thavetoinvokeit,arethegeophysicists.TheEarth'smantleissupposedtohaveaviscosityof
10
21
P.
If
younowworkout
71
2/
p,
theforceis10
41
dyn.Thatismorethan10
9
timesthegravitationalforcethathalftheEarthexertsontheotherhalf!Sotheconclu-sionis,ofcourse,thatintheflowofthemantleoftheEarththeReynoldsnumberis
very
smallindeed.Nowconsiderthingsthatmovethroughaliquid(Fig.3).TheReynoldsnumberforamanswimminginwatermightbe
10
4,
ifweputinreasonabledimensions.Foragoldfishoratinyguppyitmightgetdownto10
2.
Fortheanimalsthatwe'regoingtobetalkingabout,aswe'llseeinamo-
Copyright@1977AmericanAssociationofPhysicsTeachers
3
 
In
e
r
hoi
Force.;
visCous-fa
rces
-~1
C{.V"'f
7
a7J1_
aV-
7-
y
(~
.
C;;;o'2.
em
I.
nr
wsnr
sec
Figure
I.
ment,it'sabout
10-
4
or
10-
5.
Fortheseanimalsinertiaistotallyirrelevant.
We
knowthat
F
=
rna,
buttheycouldscarcelycareless.I'llshowyouapictureoftherealanimalsinabitbutwearegoingtobetalkingaboutobjectswhicharetheorderofamicroninsize(Fig.4).That'samicronscale,notasuture,intheanimalinFig.4.Inwaterwherethekinematicviscosityis
10-
2
em/secthesethingsmovearoundwithatypicalspeedof30/-Lm/sec.IfIhavetopushthatanimaltomoveit,andsuddenlyIstoppushing,how
7'2.
force
-
f
fo,.
wo~
7~
-4
f
10dyne!
This
f.Or-Cfa
will
towany
th/Y'l9
j
I
arJeor
J'tntAll)
af~/
n:
P
(
I
Figure2.4Am.J.Phys.,Vol.45,No.I,January1977
-
-4-
~/O
Figure3.
farwillitcoastbeforeitslowsdown?Theansweris,about0.1
A.
Andittakesitabout0.6
usee
toslowdown.IthinkthismakesitclearwhatlowReynoldsnumbermeans.In-ertiaplaysnorolewhatsoever.IfyouareatverylowReynoldsnumber,whatyouaredoingatthemomentisentirelydeterminedbytheforcesthatareexertedonyou
atthatmoment,
andbynothinginthepast.?Ithelpstoimagineunderwhatconditionsamanwouldbeswimmingat,say,thesameReynoldsnumberashisownsperm.Well,youputhiminaswimmingpoolthatisfullofmolasses,andthenyouforbidhimtomoveanypartofhisbodyfasterthan1em/min.Nowimagineyourselfinthatcondition:you'reundertheswimmingpoolinmolasses,andnowyoucanonlymovelikethehandsofaclock.Ifunderthosegroundrulesyouareabletomoveafewmetersinacoupleofweeks,youmayqualifyasalowReynoldsnumberswimmer..IwanttotalkaboutswimmingatlowReynoldsnumber
In
averygeneralway.Whatdoesitmeantoswim?Well,itmeanssimplythatyouareinsomeliquidandareallowedtodeformyourbodyinsomemanner.That'sallyoucando.
~-::=.,
I
centipoiSe
-~
~=dx/O
o
-0.1
A
J
o.~rn/crosec,
oI/stCltnce
Figure4.E.M.Purcell4
 
Figure5.
Moveitaroundandmoveitback.Ofcourse,youchoosesomekindofcyclicdeformationbecauseyouwanttokeepswimming,anditdoesn'tdoanygoodtouseamotionthatgoestozeroasymptotically.Youhavetokeepmoving.So,ingeneral,weareinterestedincyclicdeformationsofabodyonwhichtherearenoexternaltorquesorforcesexceptthoseexertedbythesurroundingfluid.InFig.5,thereisanobjectwhichhasashapeshownbythesolidline;itchangesitsshapetothedashedcontourandthenitchangesback.Whenitfinallygetsbacktoitsoriginalshape,thedottedcontour,ithasmovedoverandrotatedalittle.Ithasbeenswimming.Whenitexecutedthecycle,adisplacementresulted.Ifitrepeatsthecycle,itwill,ofcourse,effectthesamedis-placement,andintwodimensionswe'dseeitprogressingaroundacircle.Inthreedimensionsitsmostgeneraltra-jectoryisahelixconsistingoflittlekinks,eachofwhichistheresultofonecycleofshapechange.ThereisaveryfunnythingaboutmotionatlowReynoldsnumber,whichisthefollowing.Onespecialkindofswim-mingmotioniswhatIcallareciprocalmotion.Thatistosay,IchangemybodyintoacertainshapeandthenIgobacktotheoriginalshapebygoingthroughthesequenceinreverse.AtlowReynoldsnumber,everythingreversesjustfine.Time,infact,makesnodifference-onlyconfig-
Na.v/er-Stokes:
2~~;/
)'M~
-v
p
-t-
7
V
v-
=
A
iT(
If
q«/:
T/me
doesn't
matter:
The
»atrernof
moNon
is
fhe
Same,
IRhefAerflow
Or
fast
ty/'efherforwardorb~kward
/1"0
time.
The
Sea-Ilo?
Theorem
Figure6.
5
Am.J.Phys.,Vol.45,No.1,January1977
&:-
~
,;
.
~!
If
,1
r
~
Figure7.
uration.
If
Ichangequicklyorslowly,thepatternofmotionisexactlythesame.
If
youtaketheNavier-Stokesequationandthrowawaytheinertiaterms,allyouhaveleftis
\7
2
v
=
P/71,
where
P
isthepressure(Fig.6).So,iftheanimaltriestoswimbyareciprocalmotion,it
can'tgoanywhere.
Fastorslow,itexactlyretracesitstrajectoryandit'sbackwhereitstarted.Agoodexampleofthatisascallop.Youknow,ascallopopensitsshellslowlyandclosesitsshellfast,squirtingoutwater.ThemoralofthisisthatthescallopatlowReynoldsnumberisnogood.Itcan'tswimbecauseitonlyhasonehinge,andifyouhaveonlyonedegreeoffreedominconfigurationspace,youareboundtomakeareciprocalmotion.Thereisnothingelseyoucando.Thesimplestanimalthatcanswimthatwayisananimalwithtwohinges.Idon'tknowwhetheroneexistsbutFig.7showsahypotheticalone.Thisanimalislikeaboatwitharudderatbothfrontandback,andnothingelse.Thisanimalcanswim.Allithastodoisgothroughthesequencetoconfig-urationsshown,returningtotheoriginaloneatS5.Itsconfigurationspace,ofcourse,istwodimensionalwithcoordinates8
1
,8
2
Theanimalisgoingaroundaloopinthatconfigurationspace,andthatenablesittoswim.Infact,Iworkedthisoneoutjustforfunandyoucanprovefromsymmetrythatitgoesalongthedirectionshowninthefigure.Asanexerciseforthestudent,whatisitthatdis-tinguishesthatdirection?Youcaninventotheranimalsthathavenotroubleswimming.Wehadbetterbeabletoinventthem,sinceweknowtheyexist.Oneyoumightthinkoffirstasaphysicist,isatorus.Idon'tknowwhetherthereisatoroidalanimal,butwhateverotherphysiologicalproblemsitmightface,itclearlycouldswimatlowReynoldsnumber(Fig.8).Anotheranimalmightconsistoftwocellswhichwerestucktogetherandwereabletorollononeanotherbyhaving
E.M.Purcell
5

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