Three-Dimensional Flow and Lift Characteristics of A Hovering Ruby-Throated Hummingbird - Journal of The Royal Society Interface

19/09/2016 Threedimensionalflowandliftcharacteristicsofahoveringrubythroatedhummingbird|JournalofTheRoyalSocietyInterface
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Threedimensionalflowandliftcharacteristicsofa
hoveringrubythroatedhummingbird
JialeiSong,HaoxiangLuo,TysonL.Hedrick
Published9July2014.DOI:10.1098/rsif.2014.0541
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Abstract
Athreedimensionalcomputationalfluiddynamicssimulationisperformedforarubythroatedhummingbird
(Archilochuscolubris)inhoveringflight.Realisticwingkinematicsareadoptedinthenumericalmodelby
reconstructingthewingmotionfromhighspeedimagingdataofthebird.Lifthistoryandthethree
dimensionalflowpatternaroundthewinginfullstrokecyclesarecapturedinthesimulation.Significant
asymmetryisobservedforliftproductionwithinastrokecycle.Inparticular,thedownstrokegeneratesabout
2.5timesasmuchverticalforceastheupstroke,aresultthatconfirmstheestimatebasedonthe
measurementofthecirculationinapreviousexperimentalstudy.Associatedwithliftproductionisthesimilar
powerimbalancebetweenthetwohalfstrokes.Furtheranalysisshowsthatinadditiontotheangleofattack,
wingvelocityandsurfacearea,dragbasedforceandwingwakeinteractionalsocontributesignificantlyto
theliftasymmetry.Thoughthewingwakeinteractioncouldbebeneficialforliftenhancement,theisolated
strokesimulationshowsthatthisbenefitisburiedbyotheropposingeffects,e.g.presenceofdownwash.
http://rsif.royalsocietypublishing.org/content/11/98/20140541 1/27
Theleadingedgevortexisstableduringthedownstrokebutmayshedduringtheupstroke.Finally,thefull
bodysimulationresultshowsthattheeffectsofwingwinginteractionandwingbodyinteractionaresmall.
1.Introduction
Unlikebirdsoflargersizes,hummingbirdscanperformsustainedhoveringinadditiontoregularcruiseflight
andmanoeuvres.Manystudieshavebeendonetocharacterizethekinematics,physiologyand
aerodynamicsofthehummingbirdwing[14],andtheyweresummarizedintheworkofTobalskeetal.[5].
Ingeneral,hummingbirdsusesimilaraerodynamicstothoseofinsects,e.g.presenceofaleadingedge
vortex(LEV)overthewingsurface[6,7],forliftproduction.However,differencesbetweenhummingbirdand
insectaerodynamicsareconceivableastheanatomyandphysiologyofthehummingbirdwingaredistinct
fromthoseoftheinsectwing.Forexample,recentevidenceshowsthathummingbirdscanachievethe
inversionoftheangleofattackthroughactivewingrotationatthewrist[8].Thisactuationmechanismis
differentfromthatofinsectswhosewinginversioncanberealizedthroughcombinedmuscleactivationat
thewingrootandthepassivedeformationofthewingsurface[9].Theimplicationofthisdifferenceonthelift
andpowerefficiencyofhummingbirdsisstillunclear.Inordertobetterunderstandaerodynamicsof
hummingbirds,theirliftandflowcharacteristicsareneeded.Unfortunately,suchdataaresofarverylimited.
Tosetthecontextforthiswork,webrieflysummarizeseveralrecentstudiesontheforceproductionandflow
behaviourinhummingbirdflight.
Altshuleretal.[3]usedadissectedhummingbirdwingandtestedliftproductionofthewingrevolvinginone
direction.Bycomparingwithwingmodelsofincreasingrealism,i.e.thosewithsharpenedleadingedgesand
withsubstantialcamber,theyfoundthattherealhummingbirdwinggeneratesmorelift,andtheirresult
suggeststhatsomegeometricdetailssuchasthepresenceofcambertendtoincreaselift.Usingparticle
imagevelocimetry(PIV),Warricketal.[4,10]studiedtheflightofrufoushummingbirds.Theywereableto
measuretheflowinthetwodimensionalplanesthatareperpendiculartothewingaxisduringtheentire
strokecycle.BasedonthePIVdata,theyvisualizedtheLEVandcalculatedthecirculationatdifferent
spanwiselocations.Interestingly,theirresultshowsthattheaverageboundcirculationduringthedownstroke
is2.1timesofthatduringtheupstroke[10].Assumingthattheconventionalaerofoiltheoryholds,thatis,lift
islinearlyproportionaltotheboundcirculation,theauthorssuggestthattheliftproductionpossessesthe
sameamountofasymmetry.Theyfurtherproposedthepossiblemechanismsthatmayhavecontributedto
suchliftcharacteristics.Forexample,thewingvelocityandtheangleofattackduringthedownstrokeare
greaterthanthoseduringtheupstroke.Othervariablestheysuggestedincludelongerwingspanand
formationofapositivecamberduringthedownstroke.InanotherPIVstudy,Altshuleretal.[11]measured
thewakeflowofthewingsandtailofhoveringAnna'shummingbirds,andtheyproposedavortexringmodel
forthewakegeneratedbythewings.Later,Wolfetal.[12]conductedfurtherPIVstudyofthesame
hummingbirdspecies,andfromthestrengthoftheshedvortices,theyalsoconcludedthatliftproductionis
highlyasymmetricbetweenthetwohalfstrokes.
Despitethesepreviousefforts,thereexistsnodirectstudyonthedetailedforcecharacteristicsandthe
threedimensionalflowpatternofthehummingbirdwinginhoveringflight.Asausefultool,computational
fluiddynamics(CFD)hasbeenappliedinmanypreviousworkstostudyaerodynamicsofflappingwings,
includingbothrigidandflexiblewingmodels[1316].Here,wearemotivatedtoperformaCFDstudyto
quantifytheforcehistoriesinastrokecycleandtoinvestigateanyparticularforceproductionmechanisms
usedbythehummingbird.Themainquestionswewouldliketoanswerthroughthisworkinclude:(i)what
arethecharacteristicsoftheforcehistory,andwhataretheunderlyingmechanismsforthepossible
downstrokeupstrokeasymmetry?(ii)Whatisthethreedimensionalwakepatternlike,andhowmayitbe
associatedwiththeforcecharacteristics?(iii)Howmuchaerodynamicpowerdoesthehummingbirdhaveto
spendonhoveringandwhatistheefficiency?(iv)Doesthehummingbirdusesomeofthemechanismsthat
insectsuseforliftenhancement,suchaswakecaptureandwingwinginteraction?
2.Materialandmethod
2.1.Experimentandreconstructionofthewingkinematics
Thehummingbird,afemalerubythroated(Archilochuscolubris)withabodymassof3.41g,isusedasthe
subjectinthisstudy.Highspeedfilmingexperimentwasconductedtorecordthewingmotionofthebird.In
theexperiment,thebirdwastrainedtoflyina0.40.40.5m3nettedchamberandwasrecorded1000
framespersecondwitha1/5000thshutterbyfourhighspeedcameras:twoPhantomv.7.1(Vision
ResearchInc.,Wayne,NJ,USA),onePhotronSA3andonePhotron1024PCI(PhotronUSAInc.,San
Diego,CA,USA).Eachflappingcyclecontainsabout25frames.Thebirdwaslabelledpriortothe
experimentusing1mmdiameterdotsofnontoxicwhitepaint,asshowninfigure1a.Theexperimentalset
upisdescribedindetailinHedricketal.[8].Theninemarkersnumberedinfigure1aandlocatedonthe
outlineoftheleftwingareusedinthisstudy.Thesemarkersincludefivepointsontheleadingedge,oneat
thewingtipandthreeonthetrailingedge.Toavoidblindspots,thecameraswerepositionedwithone
directlybehindthebirdinthesamehorizontalplane,twowithanelevatedobliqueandslightlyrearviewand
onewithaventralviewofthebird(seetheelectronicsupplementarymaterial,figureS1).Afterthevideosare
taken,acustomMATLABprogram[17]wasusedtoautomaticallytrackthemarkersframebyframeandto
extracttheirthreedimensionalcoordinates.Aprincipalcomponentsanalysishasbeendonetoverifythat
theseninepointsaresufficienttocharacterizethewingmotion.
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Figure1.
(a)Markerpointsontheoutlineofthewing.(b)Reconstructedwingkinematics(shownfortheright
wing).(c)Thetrianglerepresentsthedistalhalfofthewingsurface,basedonwhichthechordangle,c,
andtheangleofattack,,aredefinedinthetext.(Onlineversionincolour.)
Toreconstructthewinggeometryandmotion,splineinterpolationisusedtoconnecttheoutlineofthewing
ateachinstantaneoustimeframe.Then,boththeleadingedgeandthetrailingedgeareevenlydiscretized
by41nodeseach.Thewingchordisapproximatedwithstraightsegmentswhichhaveroundedendsandan
effectivethickness7%oftheaveragechordlength.
Atriangularmeshisthengeneratedtodiscretizethewingsurface,whichisassumedtobesmooth.
Corrugationscausedbythefeathersareignoredastheireffectonthelaminarboundarylayerisexpectedto
besmallatthecurrentReynoldsnumber.DiscussionsontheeffectoffeathersathigherReynoldsnumbers
canbefoundinarecentexperimentalstudy[18].Asinglewingconsistsof1129elementsand615
Lagrangiannodes.Toincreasethetimeresolutionforthesmallstepsolutionofthesimulation,thetrajectory
ofeachmeshnodeisalsorefinedbysplineinterpolationintime.Eightcyclesofwingkinematicsare
reconstructedfromtheimagingdataandareusedforthesimulation(seetheelectronicsupplementary
material,movie).Notethatdynamicdeformationsofthewingsuchasspanwisebendingandtwistinghave
beenincludedinthereconstructedkinematics(seetheelectronicsupplementarymaterial,figureS2)and
thustheiraerodynamicconsequenceswillbeincorporatedinthesimulationresults.
Asseeninfigure1b,theentirewingsurfaceexhibitsatwistalongthewingaxis,andthetwistanglechanges
dynamicallyinastrokecycleduetothepitchingmotionofthewing.Todefinethewingpostureandthetime
varyingangleofattack,weselectthreepointsonthewing:thewingtip,theleadingedgepointandthe
trailingedgepointofthemidchord.Thesethreepointsformatriangleapproximatingposturingofthedistal
halfofthewingsurface,asindicatedinfigure1c.Thechordangle,c,isdefinedastheinstantaneousacute
anglebetweentheplanespannedbythistriangleandthehorizontalplane.Thisanglewillbeusedto
measureorientationandpitchrotationofthedistalwingsurface.Theangleofattack,,isdefinedasthe
instantaneousanglebetweenthetipvelocityvectorandthetriangle.
2.2.Simulationsetupandmodelvalidation
TheairisassumedtobegovernedbytheviscousincompressibleNavierStokesequation.Theequationis
solvedbyasecondorderaccurateimmersedboundarymethod[19]thatisabletohandlelarge
displacementofthemovingboundaries(seenumericalmethodintheelectronicsupplementarymaterial).A
fixed,nonuniform,singleblockCartesiangridisemployedtodiscretizethedomain(figure2a).The
rectangulardomainis202018cm3.Forthesinglewingsimulation,330250210(17million)points
areusedforthebaselinesimulation.Acoarsermeshwith232180140(6million)pointsandafiner
meshwith420310240(31million)pointsarealsousedinthesinglewingcasetoverifygrid
convergence.Allthreemesheshavemaximumresolutionaroundthewing,whichis0.05,0.033and0.025
Article
cminallthreedirections,forthecoarser,baselineandfinermesh,respectively.Thetwoextrasimulations
Abstract
arerunfortwocycles,andtheyproduceamaximum3%differencefromthebaselinemeshinthemeanand
1.Introduction
RMSvaluesoftheverticalforce.Thefullbodysimulationemploys336408216(30million)points,and
2.Materialandmethod
theresolutionaroundthebodyandwingsisthesameasinthebaselinecaseforthesinglewing.
3.Resultsanddiscussion
4.Conclusion
Fundingstatement
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Figure2.
Thebaselinemesharoundthebird(onlyoneoutofeveryfourpointsineachdirectionisshown).(b)Grid
convergencestudywherethenormalizedverticalforceofasinglewing,Cz,isplotted.
Thenumericalmethodhasbeenpreviouslyvalidatedforflappingwingsimulationsagainstbothexperimental
andsimulationdatainDaietal.[20],whereafruitflymodelandanimpulsivelystartedplatewerestudied.To
furthervalidatethemodelinthiswork,wecomparetheflowfieldwiththatobtainedfromthePIVexperiment
byWarricketal.[10].Notethattherufoushummingbird(Selasphorusrufus)wasusedintheexperiment,
whereastherubythroatedhummingbird(A.colubris)isusedinthisstudy.However,thesetwospeciesare
verysimilartoeachotherintermsofthemorphologicaldataandwingkinematics.Table1listssomeofthe
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keyparametersofthecurrenthummingbirdalongwiththosefromWarricketal.[10],includingthebody
mass,M,theflappingfrequency,f,thewinglength,R,thewingspan,b,averagechordlength,c,thewing
Article
area,Sandthewingbeatamplitude,.Itcanbeseenthatalltheparametersinthisstudyfallwellwithinthe
Abstract
rangesintheexperiment.Wealsoconvertedtheangleofattackandthechordangleofthepresent
1.Introduction
hummingbirdusingthedefinitionsintheexperimentalstudy,andtheresultofcomparisonisgenerally
2.Materialandmethod
consistent(inTobalskeetal.[5],thechordangleis147fordownstrokeand314forupstrokein
Warricketal.[4],theangleofattackis3612and2613.Inthisstudy,thechordangleis16for
4.Conclusion
downstrokeand48forupstrokeandtheangleofattackis33and24.Allanglesaremeasuredaccording
Fundingstatement
totheirdefinitions).TheReynoldsnumberoftheflowissettobe .Thisnondimensional
References
numberrepresentstheratiobetweenthefluidinertiaandtheviscouseffect.
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Table1. Info&Metrics Viewinline
Comparisonbetweentherubythroatedhummingbirdmodelandtheexperimentaldatafortherufous
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hummingbird.
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Figure3bshowsatypicalspanwisesliceoftheinstantaneousflowduringmiddownstrokeat70%wingspan
fromthewingroot.Notethattheexperimentaldataareshownforthesliceatapproximately80%wingspan,
or4cmfromthewingroot.Adiscussiononthechoiceofthespanwiselocationisdeferredtotheendofthis
section.Itcanbeseenthatinbothcases,astrongshearlayerexistsonthedorsalsurfaceofthewingandis
generallyattachedtothewingsurface.Intheexperiment,theshearlayerontheventralsideofthewingis
notvisibleduetotheshadoweffect.Bothfiguresshowthatalargeclockwisevortexislocatedinthewakeof
thewingandisaboutonechordlengthawayfromthetrailingedge,thoughthestrengthofvortexisweaker
inthesimulation.Overalldownwashiscreatedinbothcases,whichcorrespondstoliftproduction.Thereare
alsoothervisibledifferencesbetweenthetwoplots.Inparticular,thevorticesintheexperimentappeartobe
multipleblobsabovethewingsurface,whereasinthesimulationacontinuousvortexsheetisformedandis
slightlyseparatedfromthewingneartheleadingedge.Comparisonofothertimeframesalsodisplays
similaritiesingeneralflowpatternsbutconsiderabledifferencesinflowdetails(seetheelectronic
supplementarymaterial,figuresS3S6).Wepointoutthatvariationsinthewingkinematicsofbird
individualsmayhaveledtodiscrepanciesintheflowfieldobservedhere.Inaddition,someofthese
differencesarelikelycausedbylowresolutionintheexperimentwherearound17pointspercentimetre
wereusedforthevelocityfield.Inthesimulation,30pointspercentimetreinthebaselinegridand40points
percentimetreinthefinestgridareemployedaroundthewing.Furthermore,thetwogridsdisplayeda
consistentformofshearlayers.
Article
Abstract
1.Introduction
2.Materialandmethod
4.Conclusion
Fundingstatement
References
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Figure3.
Comparisonofthespanwisevorticity,z,duringmiddledownstroke(theunitis1s1).(a)PIVimage
adaptedfromWarricketal.[10](imageusageauthorized)(b)currentsimulation.(Onlineversionin
colour.)
Wefurthercomparetheboundcirculationaroundthewingchordwiththedatafromtheexperiment.
Figure4showsthephaseaveragedcirculation,,definedas ,alongacircularpaththat
enclosesthewingchord.Thediameterofthecircleis10%greaterthanthechordlength.Increasingthis
diameterby20%onlychangesthemaximumcirculationby5%.InWarricketal.[10],thephaseaveraged
circulationisshownat80%wingspanfortheentirestrokecycle.However,theirresultsalsoshowthatthe
spanwiselocationofthemaximumcirculationvarieslargelyamongthebirdindividuals,althoughingeneral
themaximumhappensbetween40and80%ofwingspan.Inthisstudy,wefoundthatthemaximumbound
circulationtakesplacebetween50and70%ofwingspan.Therefore,weplotfor50,70and80%wingspan
locationsandcomparethemwiththeexperimentdata.Forthesamereason,inthevalidationoftheflow
field,wechosetousethesliceat70%ofwingspan.
Article
Abstract
1.Introduction
2.Materialandmethod
4.Conclusion
Fundingstatement
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Figure4. PDF
Comparisonofthephaseaveragedboundcirculationbetweencurrentsimulationandtheexperimental
measurementofWarricketal.[10](replottedtoincludethesignof),wheretimetisnormalizedbythe
periodT.(Onlineversionincolour.)
Figure4showsthatthepresentcirculationat50%wingspanmatchesthebestwiththeexperimentaldata.
Atboth70and80%wingspan,thecirculationhasasignificantdropafterthemiddownstroke.Inthe
experimentalresult,theratioofthedownstrokeandupstrokecirculationsis2.10.1inmagnitude.Inthe
simulation,thisratiois2.2,2.3and2.0for50,70and80%wingspan,respectively.
Wefirstreporttheforces,powerandefficiencyofthesinglewingsimulationandthendiscussthe
characteristicsoftheforcesandflowfield.Intheend,wewillalsodiscussthefullbodysimulation.
3.1.Force,powerandefficiency
Theglobalcoordinatesystemisshowninfigure2a,whereX,YandZdenotetheforward,spanwiseand
verticaldirection,respectively.Theresultantforcecomponents,F X,F YandF Z,arenormalizedbythefluid
density,,theaveragewingarea,Sandtheaveragetipvelocity, ,accordingto
(3.1)
whereCX,CYandCZaretheforcecoefficientsand isthecoefficientofthesecond
momentofareaofthewingsurfaceabouttheaxispassingthroughthewingbasepointandparalleltothe
wing.Inthisstudy,S=5.68cm2and ms1areaveragedfromthereconstructedwing
motion.Theairdensityischosentobe1.23kgm 3.Fromthesedata,thereducedfrequencyofthewingas
definedby is0.16.
Article
Figure5showsthetimecoursesoftheforcecoefficientsandpowercoefficient.Notethatthecycletocycle
Abstract
variationsseeninthisfigureareduetothenonperiodicfeaturesinthewingkinematics.Theaerodynamic
1.Introduction
powerhereiscalculatedbydirectlyintegratingthedotproductbetweenthewingvelocityandthe
2.Materialandmethod
aerodynamicloadingovertheentirewingsurface.Thepowercoefficientisdefinedbynormalizingthepower
by ,where isthedimensionlessthirdmomentoftheareaofthewing.
4.Conclusion
Fundingstatement
Fromtheresult,theaverageverticalforcecoefficientis whichcorrespondsto3.12gof
References
totalweightsupportprovidedbytwowings.Thetotalliftisabout91%oftheweightofthebird.The
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remainingliftcouldbeprovidedbythewingcamber[3,4],whichisnotincorporatedintothecurrentmodel.
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Abstract
1.Introduction
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2.Materialandmethod
Figure5.
4.Conclusion
ThethreenormalizedforcecomponentsCZ(a),CXandCY(b)inthesinglewingsimulation.(c)The
powercoefficientnormalizedby Fundingstatement
,andpositivepowermeansworkdonetotheflow.
(Onlineversionincolour.) References
Figures&Data
Themoststrikingfeatureoftheverticalforceisthatthedownstrokeproducesclearlymuchhigherliftthan
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theupstroke.ThedatashowthatCZaveragedduringthedownstrokeis2.5timesofthatduringthe
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upstroke,whichisgenerallyconsistentwiththeliftestimatedbasedonthecirculationintheexperiments
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[4,10].Notethattheratiooftheboundcirculationbetweenthedownstrokeandupstrokeis2.10.1in
Warricketal.[10].Anotherobservationinfigure5isthattheforcesandpowercontainasignificantdip
duringtheupstroke.ThisdipcorrespondstotheLEVsheddingfromthewing,whichwillbediscussedin
3.4.
Theaveragedforwardforcecoefficientis ,whichismuchsmallerthan .
Theaveragespanwiseforcecoefficientis .Theseforcescanbecancelledoutforthereal
birdwhentakingintoconsiderationoftwowingsymmetry(fortheYdirection),tailmotionandpossiblythe
birdfeederinteractionintheimagingexperiment(thelattertwofortheXdirection).
Thepowercoefficientinfigure5alsoexhibitssimilarasymmetryastheverticalforcecoefficient.Further
calculationshowsthatthedownstrokerequires2.8timesasmuchpowerastheupstroke.Theaveraged
powercoefficientthroughoutthecyclesis .Definedastheratiobetweentheliftcoefficient
andthepowercoefficient,theaerodynamicefficiencyofthewingisthus .Usingthe
dimensionalvaluesof, andS,andthebodymass,weobtainthemassspecificpowerof
thebird,whichis55Wkg1.Altshuleretal.[21]estimatedthepowerconsumptionofthehummingbirds
usingtheempiricallyderiveddragcoefficientmeasuredfromarevolvinghummingbirdwing.Forthe
hummingbirdsflyingatelevationbelow1000m(bodymassrangingfrom2.5to9g),themassspecific
powerforhoveringwasestimatedtobebetween23and33Wkg1intheirwork,whichisabouthalfofthe
currentresult.
Chai&Dudley[22]reportedtheoxygenconsumptionandthereforemetabolicpowerinputofrubythroated
hummingbirdstobearound260Wkg1.Thus,ouraerodynamicpoweroutputimpliesamuscleefficiencyof
21%.Vertebratemuscleefficiencycanreachslightlylessthan30%,buthummingbirdsareexpectedtobe
slightlylessefficientbecauseofadaptationsformaintainingcontinuoushighmassspecificpoweroutputand
duetotheunmeasuredcostofacceleratingthewingmassduringeachhalfstroke.
Article
Theoverallmuscleefficiencyof21%foundhereissubstantiallygreaterthanthatreportedinearlierstudies
Abstract
[23]thatusesimplermodelstopredictaerodynamicpowerrequirementsandreportefficienciesofaround
1.Introduction
10%.However,otherrecentNavierStokessimulationsofhoveringanimalflighthavealsoreportedhigher
2.Materialandmethod
powerrequirementsthanpredicted[24]andthatrevolvingwingexperimentsdonotnecessarilyreproduce
thesameflowconditionsandthusforcecoefficientsasflappingwings[25].
4.Conclusion
3.2.Circulationandwingrotation Fundingstatement
References
Asshowninfigure4,theboundcirculationaroundthewingchordisconsistentwiththemeasurementof
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Warricketal.[10].Furthermore,thecirculationissustainedthroughthewingreversal.Forexample,during
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thedownstroke,circulationaroundthetranslatingwingisdeveloped,andtowardstheendofdownstrokeand
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throughoutsupination,thecirculationdoesnotvanishbutremainsthesamesign,e.g.positiveor
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counterclockwisefromtherightsideview.Similarly,thecirculationdevelopedduringtheupstroketranslation
remainsnegativethroughoutpronation,asshowninfigure4.Thelingeringcirculationiscausedbythe
pitchingrotationofthewingarounditsownaxis[26].Unlikeaspinningcylinderinauniformflow,this
circulationcannotalwaysbeusedforliftproduction(e.g.whenthetranslationalspeediszeroorthewing
surfaceisverticalandthushaszeroprojectedareaonthehorizontal).Therefore,theverticalforceasshown
infigure5aisstillnearlyzeroatwingreversals.
Tobetterseethephaserelationshipbetweentheliftproductionandthewingmotion,weplotinfigure6the
verticalforcecoefficient,thetranslationalvelocityofthewingasrepresentedbythetipvelocity,Vtip,the
chordanglecandalsothepitchingvelocityrepresentedby .Fewercyclesareplotted
henceforthtoshowdetailswithinacycle,althoughstatisticsaretakenfromallcyclesavailable.Fromthis
figure,wemayseeadditionalpitchingeffectotherthanpronationandsupination:duringmiddownstroke,
thereisapositivepeakin andthispeakalsoroughlycorrespondstothemaximum
translationalspeedofthewing.Suchbackwardpitchingrotationwouldincreasethecirculationand,along
withthewingtranslation,helptoenhanceliftproductionduringthedownstroke.Ontheotherhand,during
midupstroke,themagnitudeofthenegativepeakin ismuchlower.Thisdifferencecould
haveincreasedtheforceasymmetrybetweenthedownstrokeandupstroke,aswillbediscussedindetail
next.
Article
Abstract
1.Introduction
2.Materialandmethod
4.Conclusion
Fundingstatement
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Figure6.
Theliftcoefficient,thetipvelocityinmetrepersecond,thechordangleindegreesandthepitching
velocityindegreepersecondintypicalcycles(averagedatastilltakenfromalleightcycles).
3.3.Asymmetricliftproduction
3.3.1.Forceasymmetry
Figure5ashowsthatliftproductionishighlyasymmetric,withthedownstrokegeneratingmuchgreater
weightsupportthantheupstroke.Theaverageverticalforceprovidedbythedownstrokeis0.022Nandby
theupstrokeis0.0090N.Thus,theratioofasymmetryis2.5.Table2furtherliststheliftcoefficient,the
powercoefficientandthelifttopowerratioseparatelyforthedownstrokeandupstroke.Itcanbeseenthat
thedownstrokeproducesmorelift,butitisalsomorepowerconsuming.Byrescalingtheliftandpowerusing
therespectivewingvelocityandsurfaceareaofeachhalfstroketoobtain and
,weseethatthelifttopowercoefficientissimilarforthedownstrokeandupstroke.Thus,
despitethattheiraerodynamicsarequitedifferent,thetwohalfstrokesstillhavesimilarefficiency.
Table2. Viewinline
Comparisonofthedownstrokeandupstroke,where and arethelift

andpowercoefficientsrescaledbytherespective andSofeitherdownstrokeor
upstroke.
InWarricketal.[10],theforceasymmetrybetweentheupstrokeanddownstrokewasattributedtoseveral
mechanisms,includingthewingvelocity,angleofattack,wingareaandcamber.Exceptthatthecamber
effectisnotincludedinthisstudy,alltheothermechanismshavebeenobservedinthesimulation,aswillbe
discussednext.Inaddition,wefoundthatothermechanismsalsohavecontributedtotheasymmetry,which
includethedragbasedverticalforce,wingwakeinteractionandpitchingrotation.Theeffectofpitching
rotationhasbeendiscussedin3.2.Sowewillfocusontheothereffects.
Article
First,table2providesthecomparisonofafewkeykinematicparametersbetweenthedownstrokeand
Abstract
upstroke,includingtheaveragetipspeed,angleofattackandwingarea.Itcanbeseenthattheratioofthe
1.Introduction
averagewingareabetweenthedownstrokeandupstrokeisonly1.09,andtheratiooftheaveragetipspeed
2.Materialandmethod
isonly1.13.Theratiobetweenthevelocitysquaresis1.21only.Thatis,thecombinationofthewingarea
andvelocityismuchlessthantheratioof2.5intheforceasymmetry.Therefore,someothermechanisms
4.Conclusion
mustbesignificantinleadingtothelargeimbalanceoftwohalfstrokes.
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References
3.3.2.Dragbasedverticalforce
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First,weconsidertheeffectofdeviation,i.e.thenonreciprocalpathofthewinginastrokecycle.Observing
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thewingmotionfromthesideview,wenotethatthewingtiptracesaroughlyellipticalpathwhoselongaxis
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hasasmallanglewithrespecttothehorizontalplane.Thisdeviationfromthemeanstrokeplaneisshownin
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figure7abyplottingthecycleaveragedtrajectoryoftherightwingtipintheXZplane.Inthefigure,the
meanstrokeplaneistiltedforwardbyapproximately12withrespecttothehorizontal.Thisobservation
motivatesustodecomposetheforcesgeneratedbythewingintotheaerodynamiclift,i.e.theforce
perpendiculartothewingtranslation,andtheaerodynamicdrag,i.e.theforceoppositetothewing
translation,asillustratedinfigure7a.
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1.Introduction
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2.Materialandmethod
Figure7.
4.Conclusion
(a)TheaveragedtrajectoryoftherightwingtipintheXZplane.Forcedecompositionintermsofthe
aerodynamicliftanddragisillustrated.Notethattheactualdecompositiondependsontheorientationsof
Fundingstatement
thewingaxisandtipvelocityvectorinthethreedimensionalspace.(b)Theinstantaneous(thinline)and
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cycleaveraged(thickline)strokeplaneangleandangleofattackindegree.
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Todothisanalysis,wedefinetheinstantaneousstrokeplaneastheplanespannedbytheinstantaneoustip
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velocityvectorandthewingaxis.Theinstantaneousstrokeplaneangle,,istheanglebetweenthisplane
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andthehorizontalplane.Boththeinstantaneousandcycleaveragedvaluesofareplottedinfigure7b,
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alongwiththeinstantaneousandcycleaveragedangleofattack.Notethatthesetwoanglesaredefinedin
thethreedimensionalspaceandareshowninthetwodimensionalplotinfigure7aforillustrationpurposes
only.Itcanbeseenthatafterthepronation,isaround50andthendropsinmagnitudeduringmorethan
halfofthedownstroke.Duringearlydownstroke,theangleofattackislargeanddropsfrom80to39.The
twoanglehistoriesindicatethatduringearlydownstroke,thewingispressingdownwardwhilesweeping
forward.Towardstheendofdownstroke,becomespositive,butitsmagnitudeislessthan25before
supination.Incomparison,duringtheupstrokeisaround10andonlyvariesslightly.
Wedefinetheresultantforcenormaltotheinstantaneousstrokeplaneaslift,F L,andtheforceoppositeto
thedirectionoftheinstantaneoustipvelocityasdrag,F D.Figure8showsthenormalizedliftanddragby
,CLandCD,andalsotheirprojectionsintheverticaldirection,CL,ZandCD,Z,fortwo
cycles.Infigure8a,CLandCDcorrelatewitheachotherandhavesimilarmagnitude.Theaveragedataare
listedintable3separatelyforthedownstrokeandupstroke.Figure8bshowsthatduringthedownstroke,
draghasasignificantpositivecontributiontotheverticalforceduringthefirsthalfofthedownstrokeandhas
onlyasmallnegativecontributionduringthesecondhalfofthedownstroke.Duringtheupstroke,draghasa
mostlynegativecontribution,andthemagnitudeissmall.Onaverage,thedragbasedverticalforce,CD,Z,is
0.63or24%ofthetotalverticalforceCZduringthedownstroke,anditis0.12or12%ofCZduringthe
upstroke.AsCD,Zofthedownstrokeis61%ofCZoftheupstrokeandCD,Zoftheupstrokeissmall,wecan
concludethatdragcontributes0.61outoftheasymmetryratio2.5intheverticalforce.
Table3. Viewinline
Theaerodynamiclift,dragandtheirverticalcomponentforbothdownstrokeandupstroke.Allforcesare
normalizedinthesamewayasdescribedearlier.
Article
Abstract
1.Introduction
2.Materialandmethod
4.Conclusion
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Figure8.
(a)Normalizedaerodynamicliftanddragfortwotypicalcycles.(b)VerticalcomponentsofCLandCD.
Averagedataarecalculatedfromalleightcycles.(Onlineversionincolour.)
Figure8a,balsoshowthatafterexcludingthedragbasedverticalforce,theliftcoefficient,CL,isstill
asymmetricbetweenthedownstrokeandupstroke,andsoisitsverticalcomponent,CL,Z.Onaverage,the
downstroketoupstrokeratioinCLis1.80.
3.3.3.Wingspeedandangleofattack
AspointedoutbyWarricketal.[4],thedifferencesinthetranslationalspeedandangleofattackbetween
thedownstrokeandupstrokemayhavebeenamajoreffectfortheliftasymmetry.Totestthishypothesis,we
designedarevolvingwingmodelforthecurrenthummingbird.Inthismodel,arigidwingwithaflatsurfaceis
createdbyprojectingtheactualwingduringamiddownstrokeontoaplane(sothespanwisetwistis
eliminated),andthemodifiedwingacceleratesfromthestationarypositiontoamaximumvelocityandthen
continuestorevolveatthatvelocity(seetheelectronicsupplementarymaterial,figureS7).Thestrokeplane
ishorizontal,andtheangleofattackiskeptconstantthroughouttheentireprocess.Twocasesare
simulatedinthistest.Inthefirstcase,thewingtipfollowsthetranslationhistoryofthewingtipinanactual
downstrokechosenfromonetypicalcycle,from0tothemaximumvelocity15ms1withinthetimeperiod
0.2T,andtheangleofattackis=41.Inthesecondcase,thewingtipfollowsthetranslationhistoryofan
actualupstrokeofthesamecycle,from0tothemaximumvelocity12ms1withinthetimeperiod0.15T,
andtheangleofattackis=28.Theairproperties(densityandviscosity)remainthesameinthissetup.
Theresultsshowthattheratiooftheliftduringsteadytranslationis1.57betweenthedownstrokerevolving
wingandtheupstrokerevolvingwing.Thus,thecombinedeffectoftranslationandangleofattackis
Article
confirmed.However,itshouldbenotedthatcomparingtherevolvingwingandflappingwing,thetransient
Abstract
historiesofliftdisplayconsiderabledifferences,asseeninfigure9.Thisresultsuggeststhattherotational
1.Introduction
motionoftheflappingwingduringtheaccelerationphaseisstillimportant.
2.Materialandmethod
4.Conclusion
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Figure9.
Therevolvingwingversustheflappingwingintheproductionoflift.(a)Downstrokeand(b)upstroke.
Onlyonetypicalstrokecycleisusedinthistest.Theverticaldashedlinerepresentsthetimeperiodof
wingacceleration.Allforcesarenormalizedbythesamefactor.(Onlineversionincolour.)
3.3.4.Wingwakeinteraction
Wingwakeinteractionisauniquefeatureofflappingwings.Inthepreviousstudyoftheaerodynamicsofthe
fruitfly,Dickinsonetal.[26]suggestthatthewingwakeinteractionenhancesliftproductionandisableto
generateapeakforceatthebeginningofahalfstrokeiftheangleofattackisreversedtimely(whichisthe
caseforadvancedpitchingandsymmetricpitching).Itwouldbeinterestingtoseetowhatextentasimilar
effectexistsinhummingbirdflight,andalsowhetherthiseffectinfluencesthedownstrokeandupstroke
differently.
First,theliftgraphinfigure5showsthatthereisnoclearpeakinCZinthebeginningofeitherdownstroke
orupstroke.Toinvestigatethepresenceofthewingwakeinteraction,infigure10wevisualizetheflowina
Article
XZplaneshortlyafterthewingreversalbyplottingthevelocityvectorstangenttotheplane.Infigure10a
Abstract
whereatypicaldownstrokeisshown,thewingmovessomewhatdownwardandtranslatesatalower
1.Introduction
elevation,andthisallowsthewingtocapturetheoppositeflowproducedbytheprecedingupstroke.Note
2.Materialandmethod
thattheoppositeflowalsotravelsdownwardduetotheoveralldownwash.Ontheotherhand,infigure10b
whereatypicalupstrokeisshown,thewingmovessomewhatupwardandtranslatesatahigherelevation,
4.Conclusion
andthusitmissestheoppositeflowproducedbytheprecedingdownstroke.Therefore,qualitatively
Fundingstatement
speaking,thedownstrokebenefitsmorefromthewingwakeinteractionthantheupstroke,althoughthe
References
interactiondoesnotgenerateaseparateliftpeakbecauseofitstiming.
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PDF
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Article
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Abstract
1.Introduction
Downloadpowerpoint
2.Materialandmethod
Figure10.
4.Conclusion
Typicalasymmetricwingwakeinteractionshowninaspanwiseslicefor(a)anearlydownstrokeatY=
70%wingspanandt/T=0.1,and(b)anearlyupstrokeatY=50%wingspanandt/T=0.6.Thedashed
Fundingstatement
lineindicatesthetiptrajectoryofthiscycle,andthecircleindicatestheoppositeflowcausedbythe
References
precedinghalfstroke.Thechordisshownasathickline.Thebirdbodywasnotincludedintheactual
singlewingsimulation.(Onlineversionincolour.)
Figures&Data
Info&Metrics
Tofurthertheinvestigation,wesimulateeachhalfstrokeinseparaterunswithotherwiseidenticalwing
eLetters
kinematics.Thestartandendofthesimulationarebasedontheobservationofthewingpositionsat
PDF
pronationandsupination.Thus,theeffectofwingwakeinteractionisexcludedinsuchisolatedwingstrokes.
Oneissuetobearinmindisthatintheisolatedstrokes,thewingdoesnotencounterameandownwashasit
doesinthecontinuousstrokes.Thedownwashreducestheeffectiveangleofattackandthusweakenslift
production.
Figure11showstheliftcoefficient,CZ,oftheisolatedstrokesalongwiththedataforthecontinuousstrokes.
Inthefirstdownstroke,thetwosimulationsproduceidenticalresultsandthusarenotshown.Fortheother
strokes,notabledifferencescanbeseenbetweenthetwosimulations.Fordownstrokes,liftproducedbythe
isolatedstrokesisclosetothatproducedbythecorrespondingcontinuousstrokes,whereasforthe
upstrokes,theisolatedstrokesproducegreaterliftthanthecontinuousstrokes.Onaverage,theratiooflift
betweenthecontinuousandtheisolatedstrokesis93.2%fordownstrokeand83.1%forupstroke.Thisresult
suggeststhatforthepresenthummingbird,theliftenhancingeffectofthewingwakeinteractiondoesnot
exceedthemitigatingeffectsofotherpossiblemechanismspresent,e.g.thedownwash.Ontheotherhand,
thewingwakeinteractiondoesaffecttheliftasymmetry,asthedownstroketoupstrokeratiointhevertical
forceisreducedto2.2fortheisolatedstrokes.
Article
Abstract
1.Introduction
2.Materialandmethod
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4.Conclusion
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Openinnewtab
References
Figures&Data
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Figure11.
eLetters
Liftproductionofisolatedwingstrokesandcontinuousstrokes.Averagedataarecalculatedfromalleight
cycles.(Onlineversionincolour.)
PDF
Finally,itshouldbenotedthattheupstrokedownstrokeforceasymmetrywasalsoobservedinthehovering
flightofsomeinsectssuchasthehawkmoth[24]andfruitfly[27],thoughforthefruitflytheupstroke
producesgreaterverticalforce.Itmaybepossiblethatsomeoftheeffectsdiscussedinthisstudyhaveled
totheobservedasymmetry.Forexample,fromthetiptrajectoryofthoseinsectwingsandtheforcehistory
providedintheearlierstudies[24,27],onecanseeasimilarcorrespondencebetweenthedownwardwing
translationandthelargeliftproduction,whichislikelycausedbyasimilardragbasedeffect.
3.4.Threedimensionalvortexstructures
Figure12showsafewselectedsnapshotsofthethreedimensionalflowfieldinastrokecycle,whichis
identifiedbyplottinganisosurfaceoftheimaginarypartofthecomplexeigenvalueoftheinstantaneous
velocitygradienttensor[28].Thistechniqueallowsonetoidentifyregionswhererotationdominatesover
strain.
Article
Abstract
1.Introduction
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2.Materialandmethod
Openinnewtab
4.Conclusion
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Downloadpowerpoint
References
Figure12.
Figures&Data
Threedimensionalvortexstructuresintheflowduringastrokecycle,wherethetimestampfrom(a)to
Info&Metrics
(d)is0.37,0.51,0.58and0.78T.Thevortexloopfromthedownstrokeismarkedbyadashedline.The
thickarrowin(d)indicatesthelocationwheretheLEVispinchedoff.(Onlineversionincolour.)
eLetters
PDF
AnLEVisdevelopedinthebeginningofthedownstroke,andthisLEVgrowsstrongerandremainsstably
attachedtothewingduringmostofthedownstroke.Duringwingtranslation,theLEV,thetipvortex(TV)and
theshedtrailingedgevortex(TEV)areconnectedendtoend,formingavortexloop,withinwhichtheair
movesdownward(figure12a).Towardstheendofdownstroke,thewingrotatesrapidlyalongitsownaxis,
andtheLEVisdividedintotwobranches,knownasdualLEV[29],asseeninfigure12a.Correspondingto
thestableLEV,thereisnoclearliftdropthroughoutthedownstroketranslation.Attheendofdownstroke,
theLEVstartstoshedfromthewingasseeninfigure12b.Anotherfeatureofthedownstrokeisthatthe
wingcatchesthevortexloopproducedbytheprecedingupstrokeanddisruptsthisloopthroughthewing
wakeinteraction.
Duringtheupstroke,anLEVisalsoformedinthebeginning(figure12c),butthedistalportionofthisLEVis
pinchedoffduringmidupstroke,asshowninfigure12d.Correspondingly,thereisavisibledipinthevertical
forcearoundthesametimeoftheupstrokeshowninfigure5a.Later,theLEVwillbeformedagainandwill
alsoformbrancheslikedualLEVs.Asdiscussedearlier,duringtheupstrokethewingmissesthewake
producedbytheprecedingdownstroke.Asaresult,thevorticesgeneratedbythedownstrokearebetter
preservedinthewake.
Figure12alsoshowsthatthewakecontainsmanyslendershapedvortices.Thesevorticesareformed
mainlyduetobreakupoftheTVandTEVatthecurrenthighReynoldsnumber.Thisflowbehaviouris
consistentwiththeresultofarecentwork[29]thatdemonstratedasimilarphenomenonofvortexbreakup
atRe=1500.Tofurtherconfirmtheaccuracyofthesevortices,wehavecomparedthesimulationsfromthe
baselineandthefinestmeshasdescribedin2.2,andtheresultsshowthatthegeneralcharacteristicsof
thevorticesareconsistent.Intheregionsfarawayfromthewing,theisolatedvorticeslikelycontainartificial
effectsduetoreducedresolutionthere.
3.5.Fullbodysimulation
Afullbodymodelofthehoveringhummingbirdisalsocreatedbyusingsymmetrickinematicsfortheleftand
rightwings.Thebodyofthebirdisapproximatedbyasequenceofellipseswithdifferentsizesandaspect
Article
ratios.ThebirdmodelisruninanextendeddomainintheYdirection.Thetypicalflowfieldisshownin
Abstract
figure13formiddownstrokeandshortlyaftersupination.Fromthevortexstructuresintheflow,wenote
1.Introduction
thatLEVandtheTVduringthedownstrokearesimilartothoseinthesinglewingsimulation.However,
2.Materialandmethod
duringsupination,thetwowingsareneareachother(theincludedangleisabout30).Theflowsaroundthe
twowingsarecloseenoughtointeract.Inparticular,whenthewingsmoveawayfromeachother,the
4.Conclusion
vorticesgeneratedfromeachwinggrowandcollidewithoneanother.Theinteractioniscomplexandleads
Fundingstatement
tofurtherbreakupofthevortices(seeamovieintheelectronicsupplementarymaterial).Otherthanthat,
References
themajorvortexstructures,suchastheLEVandTV,aresimilartothoseseeninthesinglewingsimulation.
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Article
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1.Introduction
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2.Materialandmethod
Figure13.
4.Conclusion
Threedimensionalvortexstructuresinthefullbodysimulationshownforadownstrokeat0.37T.(Online
versionincolour.) Fundingstatement
References
Despitetheeffectofthewingwinginteractiononthethreedimensionalvortexstructures,theliftproduction
Figures&Data
isnotsignificantlyaffected.Figure14providesacomparisonoftheliftcoefficientbetweenthefullbodyand
Info&Metrics
thesinglewingsimulations.Itcanbeseenthattheforcesfromthetwosimulationsareveryclosetoeach
eLetters
other.Thisresultsuggeststhatthewingwinginteractionandthewingbodyinteractiondonotplayan

importantroleinliftproductionofthehummingbird.
PDF
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Article
Openinnewtab
Abstract
1.Introduction
Downloadpowerpoint
2.Materialandmethod
Figure14.
4.Conclusion
Comparisonoftheverticalforcecoefficientintypicalcyclesbetweenthefullbodysimulationandsingle
wingsimulation.(Onlineversionincolour.)
Fundingstatement
References
Figures&Data
4.Conclusion
Info&Metrics
eLetters
Athreedimensionalsimulationwasperformedforahoveringhummingbirdwiththerealisticwingmotion
reconstructedfromimagingdata.Thesimulationcapturestheliftandpowercharacteristicsinastrokecycle
PDF
andalsodetailsoftheflowfield.Ourresultconfirmsandprovidesspecificdatafortheliftasymmetrythat
waspreviouslysuggestedbasedonthemeasurementofthecirculationaroundthewing.Furthermore,we
quantitativelyanalysedthesourcesoftheliftasymmetryandpointedoutthemechanismsthatleadtothe
asymmetry.Summarizingtheresults,thedownstrokeproduces150%higherverticalforcethantheupstroke.
Amongthefactors,thewingareacontributes10%greaterforce,thedragbasedeffectcontributes60%,the
wingwakeinteractioncontributes30%andtheremaining50%canbeattributedtothecombinedwing
speed,angleofattackandwingrotation.
Fundingstatement
ThisresearchwassupportedbytheNSFunderCBET0954381toH.L.andIOS0920358toT.L.H.The
computingresourceswereprovidedbytheNSFXSEDEandtheVanderbiltACCRE.
ReceivedMay21,2014.
AcceptedJune18,2014.
2014TheAuthor(s)PublishedbytheRoyalSociety.Allrightsreserved.
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Three-Dimensional Flow and Lift Characteristics of A Hovering Ruby-Throated Hummingbird - Journal of The Royal Society Interface

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