Three-Dimensional Simulation For Fast Forward Flight of A Calliope Hummingbird - Open Science

19/09/2016 Threedimensionalsimulationforfastforwardflightofacalliopehummingbird|OpenScience
Institution:FundaoUniversidadedoRioGrande

search
Advanced
Home Content Informationfor Aboutus Signup Submit
Threedimensionalsimulationforfastforward
flightofacalliopehummingbird
JialeiSong,BretW.Tobalske,DonaldR.Powers,TysonL.Hedrick,
HaoxiangLuo
Published8June2016.DOI:10.1098/rsos.160230
Article Figures&Data Info&Metrics PDF
Abstract
Wepresentacomputationalstudyofflappingwingaerodynamicsofacalliopehummingbird(Selasphorus
calliope)duringfastforwardflight.Threedimensionalwingkinematicswereincorporatedintothemodelby
extractingtimedependentwingpositionfromhighspeedvideosofthebirdflyinginawindtunnelat8.3ms
1.Theadvanceratio,i.e.theratiobetweenflightspeedandaveragewingtipspeed,isaroundone.An
immersedboundarymethodwasusedtosimulateflowaroundthewingsandbirdbody.Theresultshows
thatbothdownstrokeandupstrokeinawingbeatcycleproducesignificantthrustforthebirdtoovercome
dragonthebody,andsuchthrustproductioncomesatpriceofnegativeliftinducedduringupstroke.This
featuremightbesharedwithbats,whilebeingdistinctfrominsectsandotherbirds,includingcloselyrelated
swifts.
1.Introduction
http://rsos.royalsocietypublishing.org/content/3/6/160230 1/27
Hummingbirdsaredistinguishedandextremelyagileflyersamongbirds.Theyarecapableofnotonly
sustainedhoveringflight,butalsofastforwardflightandvariousrapidmanoeuvers.Recentstudiesoffluid
dynamicshavemainlyfocusedonhummingbirdsuniquehoveringcapabilityandunsteadyaerodynamics
associatedwiththewingsthatmoveinarelativelyhorizontalplane[16].Thesestudiesincludeboth
experimentalmeasurementusingdigitalparticleimagevelocimetry(PIV)[14]andcomputationalfluid
dynamicssimulations[5,6].Despitehavingavertebratemusculoskeletalconfiguration,hummingbirdsin
hoveringmovetheirwingsforwardduringdownstrokeandbackwardwithaninvertedangleofattackduring
upstroke,i.e.aflightstrategythatisusedbymanyinsects(e.g.fruitfliesandbumblebees)aswellasbats.
Asaresult,bothdownstrokeandupstrokeofthewingsgenerateweightsupport,eventhoughithasbeen
shownforsomeofthehummingbirdsthatdownstrokeproducesapproximatelytwiceasmuchasweight
supportasupstroke[2,5].Closelyassociatedwiththetranslationandpitchingmotionsofthewings,the
unsteadyflowaroundhummingbirdshassimilarfeaturesasthoseoftheinsectwings[710].Forexample,
theflowislargelyseparatedfromthetopwingsurfaceleadingedgevorticesareformednearthewing
surfaceduringbothdownstrokeandupstroke,andtheyplayanimportantroleintheforceproduction.From
abiologicalperspective,suchsimilaritywithinsectsappearstobeowing,inpart,toevolvedspecializationfor
hightransmissionratio(i.e.theratioofcontractilevelocityofthemuscletotangentialvelocityofthewingtip)
inthehummingbirdwing[11].
Comparedwithhoveringflight,theforwardflightofhummingbirdsanditsfluiddynamicshavebeenmuch
lessstudied.Tobalskeetal.[12]performedcomprehensivemeasurementoftheflightkinematicsofthe
rufoushummingbirdsinawindtunnelatspeedsrangingfromzero(hovering)to12ms1.Thedatathey
obtainedincludethebodyorientationangle,wingbeatfrequency,wingbeatamplitude,strokeplaneangle,
wingtiptrajectoryandtimedependentvariablessuchasthewingchordangleandwingarea.Ingeneral,as
theflightspeedincreases,thebirdsaligntheirbodiesmoreparalleltotheflowtoreducedrag,andthe
strokeplanebecomesmorevertical,whichisbeneficialforthrustproduction.Basedontheirdata,the
advanceratio,J,definedastheratiobetweentheflightspeed,U,andtheaveragewingtipspeed,Utip,is
betweenJ=0forhoveringandJ=1.5forthemaximumspeedat12ms1.Incomparison,insectstypically
haveanadvanceratioofJ<1[13,14].Forexample,fruitfliesandbumblebeeshavetheadvanceratioat0.25
and0.6,respectively[15,14]sotheirwingspeedismuchfasterthantheflightspeed.Suchadifferencein
theflightdynamicsimpliesthatthereshouldexistasignificantlydifferentforceproductionmechanisminthe
hummingbirdwingsthanthatofthoseinsectwingsduringforwardflight.
Amonganimalflyers,thereareseveraltypesofthrustproductionmechanismsfortheforwardflightmode.
Onecommontypeisthesocalledbackwardflick[16],whichisusedbymanyinsects,e.g.
bumblebees[14,15,17],alsobybats[18]andbirdsthathaverelativelypointedwings(e.g.doves)[19]during
slowflightwithJ<1.Inthistype,thestrokeplanetiltsforwardandthebackwardspeedofthewingduring
upstrokeisfasterthantheflightspeed.Thus,theaerodynamicliftgeneratedbythewingsisdirectedforward
duringupstrokeandfunctionsasthrust.Anothertypeisthepaddlingmodefoundinarecentstudyoffruit
fliesatspeedof0.43ms1[20].TheadvanceratiooffruitfliesisnearJ=0.25[14].Inthiscase,thestroke
planeoftheinsectsremainsnearlyhorizontal,andtheangleofattackofthewingsatupstrokeismuch
greaterthanthatatdownstroke.Thus,largedragisproducedintheforwarddirectionasadragbasedthrust
mechanism.Forlargebirdsatcruisingflight,theadvanceratioisusuallyaboveone[21],andthrustis
typicallygeneratedduringdownstrokewhentheleadingedgetiltsdownwardtoredirecttheaerodynamiclift
forwardforsimultaneousweightsupportandthrustproduction[22]duringupstroke,birdswithrounded
wingstendtofullyflexandfeathertheirwings,thusproducinglittleforce[2225],butthosewithpointed
wingssuchaspigeons,doves,cockatielsandparakeetssweeptheirwingsonupstrokeinfastflightand
continuetoproduceweightsupport,albeitatareducedlevelcomparedwithdownstroke[26,27].
Sincetheadvanceratioofhummingbirdsvariesfromzerotoaboveone,itispossiblethattheyusedisparate
forceproductionmechanismsatdifferentflightspeeds.FromTobalskeetal.[12],thetiptrajectoryofthe
hummingbirdsatslowflightspeedsishighlyskewed,whenviewedfromaglobalcoordinatesystem,and
resemblesthatofinsects.Thus,thebackwardflickmodeisprobablyemployedforthrustproduction.
However,atfastspeeds,itisnotclearwhetherthehummingbirdsbecomemorelikeotherbirds,orifthey
adoptadifferentflightstrategy.Toanswerthisquestion,itisnecessarytoexaminethedetailedwingmotion
atthosespeeds.Inadditiontothekinematicanalysis,accuratecalculationoftheforcesisneeded,asthe
flowunderconsiderationishighlythreedimensionalandinvolvesunsteadyeffectsbeyondlimitationsof
quasisteadymodels.
Inthecurrentstudy,weaimtounderstandliftandthrustproductionofhummingbirdsduringfastforward
flight.Theflowfieldandbehaviourofvorticeswillbeinvestigatedalongwithaerodynamicforces.Inaddition,
wewillcompareforcegenerationmechanismsbetweenhummingbirdsandinsects,otherbirdsincludingthe
closestrelatives,swifts,aswellasbats.Followingtheapproachinapreviousstudyofhoveringflightofthe
hummingbird[5],wedevelopahighfidelitycomputationalmodelthatincorporatestherealistickinematicsof
thebirdwingsandadoptthreedimensionalnumericalsimulationstoresolvetheunsteadyflow.
2.Modelconfigurationandsimulationapproach
2.1Reconstructionofthewingkinematics
Afemalecalliopehummingbird(Selasphoruscalliope)wasthestudysubject,whosebasicmorphological
dataareprovidedintable1.Theexperimentalstudywasconductedtoobtainthewingkinematicsata
sustainedflightspeedofU=8.3ms1,atwhichthewingbeatfrequencyis45.5Hzwhilethestrokeplane
anglebetweenthestrokeplaneandthehorizontalis67.9.Intheexperiment,thebirdwasplacedinan
opencircuit,variablespeedwindtunnelwithafeederatthemiddleofthetunnel,anditwastrainedtoadapt
tothewindwhilefeeding.Werecordedflightkinematicsofthehummingbirdusingthreehighspeedcameras
distributedoutsidethewindtunnel:1PhotronSA3(PhotronUSAInc.,SanDiego,CA,USA)and2two
Photron1024PCI,allwithelectronicallysynchronizedshuttertiming.Twocameraswereplaceddorsallyto
thebird,andonewasplacedlaterally,asshownbytheviewsinfigure1.Videorecordingsweremadeat
1000Hzwithashutterspeedof1/10000s.Theworkingsectionofthetunnelis85cminlength,squarein
crosssection,6060cm2attheinletandincreasingto61.561.5cm2attheoutlettoaccommodate
boundarylayerthickening[28].Maximumdeviationsinvelocitywithinacrosssectionarelessthan10%of
themeantheboundarylayerislessthan1cmthickandturbulenceis1.2%.
Downloadfigure
Openinnewtab
Downloadpowerpoint
Figure1.
Cameraviewsofthehummingbirdflyinginthewindtunnel.
Table1. Viewinline
Morphologicalandbasickinematicsdataofthehummingbirdusedinthestudy.
Afterthevideosweretaken,acustomMATLABprogram[29]wasusedtotrackthemarkersframebyframe
andtoextracttheirthreedimensionalcoordinates.Thesemarkerswereprelabelledonthewingsusingnon
toxicpaint,andtheyincludedfivepointsontheleadingedge,onatthewingtipandthreeonthetrailing
edge,asshowninfigure2,whereacomparisonofthereconstructedmodelwiththecameraviewshows
thattheinstantaneouswingpositionanddeformationarewellcaptured.Inasimilarstudyforthehovering
hummingbird[5],aprincipalcomponentsanalysisverifiesthatthesepointsaresufficienttocharacterizethe
wingmotion.Thewinggeometryreconstructionprocessissimilartothatinthepreviousstudy[5].Thewing
profilewasconstructedusingsplineinterpolationthroughthemarkerpoints,andthewingsurfacewasthen
builtusingtriangularelementswithintheprofile.Thebirdbodywasreconstructedusingthecameraviewsof
thehummingbird.Inthecurrentreconstruction,asinglewingconsistsof1335elementsand718nodes,
whilethebodysurfaceconsistsof3560elementsand1782nodes.Atotalof13cyclesofwingbeatsduring
steadyflightwerecaptured,andeachcyclecontainsapproximated22frames.Toincreasethetime
resolutionofthewingposition,thetrajectoriesofthewingmeshnodesarealsorefinedbyspline
interpolationintime.Sevencyclesofwingbeatswerereconstructedfromtheimagingdataandusedforthe
simulation.Figure3showsasequenceofwingpositionswithinacycleandalsothewingtiptrajectory(see
theelectronicsupplementarymaterialforananimation).
Downloadfigure
Openinnewtab
Downloadpowerpoint
Article
Figure2.
Abstract
Reconstructedwingpositionsandcorrespondingsnapshotsfromthecameraview.
1.Introduction
3.Resultsanddiscussion
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Figures&Data
Info&Metrics Downloadfigure
PDF
Openinnewtab
Downloadpowerpoint
Figure3.
Reconstructedwingpositionofthehummingbirdwithinonebeatcycle:(a)downstroke,(b)upstrokeand
(c)theleftwingtiptrajectoryasviewedinabodyfixedcoordinatesystem,wherethethicklineisthe
phaseaveragedandthethindashedlineistheinstantaneousdata.
2.2Wingkinematics
Figure4showsboththeinstantaneousandphaseaveragedwingtipvelocityofthehummingbird.The
upstrokehasslightlyhighervelocitythandownstroke.Thepeakvalueisat11.22ms1fordownstrokeand
12.18ms1forupstroke.Thewingtipvelocityaveragedthroughoutthecyclesis8.14ms1,whichgivesthe
advanceratioJ=1.02.Thewingareacanbecalculatedfromthereconstructedkinematicsanditvaries
between5.34cm2duringdownstrokeand5.03cm2duringupstroke,withaverageatS=5.18cm2.
We use cookies to help us improve this website. Learn more Close
Downloadfigure
Article
Abstract
Openinnewtab
1.Introduction
Downloadpowerpoint
Figure4.
4.Conclusion
Instantaneous(thindashedlines)andaverage(thickline)wingtipvelocity.Inallfigures,greybackground
Ethics
representsdownstrokeandwhiterepresentsupstroke.
Dataaccessibility
Competinginterests
Fromfigure3,wingtwistalongtheaxisandspanwisebendinginacycleareevident.Tocharacterizethe
Funding
positionofacrosssectionofthewings,wedefinethechordangleastheinstantaneousanglebetweenthe
Acknowledgements
chordandtheflightdirection.Theangleofattack,,isdefinedastheanglebetweenthechordandthe
References
relativeflowdirectionthatcombinesboththefreestreamvelocityandthetranslationalvelocityofthechordat
Leaveacomment
theleadingedge.Thesetwoanglesareplottedinfigure5fortwochordsandfivecycles,oneproximal
Figures&Data
chordatdimensionlesslocationr^ = r/R = 0.15 andonedistalchordatr^ = 0.9 ,whicharedenotedby
Info&Metrics
subscriptspandd,respectively.
PDF
Downloadfigure

Openinnewtab
Article Downloadpowerpoint
Abstract
Figure5.
1.Introduction
Chordangle(a,b)andeffectiveangleofattack(c,d)foraproximalchordatr^ = 0.15 (a,c)anda
distalchordat ^ = 0.90 (b,d).
r
Fromtheseplots,wecanseelargedifferencesbetweentheproximalchordanddistalchord.Forthe
4.Conclusion
proximalchord,thechordangle
Ethics pandangleofattackparebothpositiveduringtheentirecycle.Forthe
distalchord,theseangleschangethesignandvarysignificantly.Duringthedownstroke,
Dataaccessibility disnegative,i.e.
theleadingedgetiltingdownward,but
Competinginterests dispositiveowingtofasttranslationofthechord.Duringupstroke,
dispositive,i.e.theleadingedgetiltingupward,butthe
Funding disnegative,indicatingthatthepressuresurface
Acknowledgements
andthesuctionsurfaceareswappedatthatmoment.Wingtwistcanbedescribedbythedifferencebetween
References
thetwochordangles,dp,whichisplottedinfigure6.Itisshownthatthetwistanglereachesitsextreme
Leaveacomment
valueduringmiddownstrokeandmidupstrokehowever,itismorepronouncedduringupstroke(near40)
Figures&Data
thanduringdownstroke(near25).Thesedifferencesbetweentheproximalsectionandthedistalsection
Info&Metrics
leadtohighlynonuniformpressuredistributiononthewingsurfaceasshownlater.
PDF
Downloadfigure
Article
Abstract Openinnewtab
1.Introduction
Downloadpowerpoint
Figure6.
4.Conclusion
Wingtwistasmeasuredusingthedifferencebetweentwochordangles.
Ethics
Dataaccessibility
2.3Simulationsetupandverification
Competinginterests
Inthemodel,theReynoldsnumber,definedasU c
/ ,issettobeRe=3000,where c
istheaveragechord
Funding
lengthandisthekinematicviscosity.Theflowisassumedtobegovernedbytheviscousincompressible
Acknowledgements
NavierStokesequation,whichissolvedbyaninhousecodethatadoptsasecondorderimmersed
References
boundaryfinitedifferencemethod.Thecodeisabletohandlelargedisplacementofthemoving
Leaveacomment
boundaries[30].Afixed,nonuniform,singleblockCartesiangridisemployedtodiscretizethedomain
Figures&Data
(figure7a).Therectangulardomainis252016cm3.Forthesimulation,704842560(333million)points
Info&Metrics
areusedforthebaselinesimulation.Afinermeshisalsousedinthesimulationtoverifygridconvergence.
PDF
Bothofthesemesheshavemaximumresolutionaroundthewing,whichis 60
1
cminallthreedirectionsfor
thebaselinecaseand 1
cm fortherefinedcase.Thesimulationwasruninparallelusingdomain
70
decompositionandMessagePassingInterface(MPI).Thetimestepist=5s,whichleadstoapproximately
4400stepsperwingbeatcycle.AmultigridmethodwasemployedtoaccelerateconvergenceofthePoisson
solver.Atotalof96processorcoreswereusedforthebaselinecase,and128coresfortherefinedcase.
Downloadfigure
Article
Abstract Openinnewtab
1.Introduction
Downloadpowerpoint
Figure7.
4.Conclusion
(a)Baselinemesharoundthebird(onlyoneoutofevery12pointsineachdirectionisshown).(b)Force
comparisonbetweenbaselinesimulationandfinemeshsimulation.
Ethics
Dataaccessibility
TheverticalforceF ZandthrustF T=F Xgeneratedbyonewingarenormalizedbythefluiddensity,,the
Competinginterests
flightspeed,U,andthesurfaceareaofthewingaccordingto
Funding
Acknowledgements FZ FT
CZ = and CT = .
References (1/2)U
2
S (1/2)U
2
S
Leaveacomment
(2.1)
Figures&Data
Theliftanddragonthebirdbody,F Z andF D ,arenormalizedinthesamemanner.Theaerodynamicpower
Info&Metrics b b
coefficientofonewingisdefinedas
PDF
f u dA
CP = .
3
(1/2)U S
(2.2)
wheref isthestressonthewingsurface,andu isthevelocityofapointonthewinginthebodyfixed

coordinatesystem.
Thesimulationresultsfortwowingbeatcyclesfrombothmeshesareshowninfigure7andtable2for
comparison.Infigure7andalsootherfiguresfromherein,theshadedareaindicatesdownstroke,whilethe
whiteareaindicatesupstroke.Theseresultsincludethetimeaveragedliftandthrustofonewingandalsolift
anddragofthebirdbody.Fromthetable,weseethatthemaximumdifferenceofalltheforcesislessthan
5%.Thus,thebaselineresolutionisdeemedsatisfactoryforthecurrentstudy.
Table2. Viewinline
Comparisonoftheforcecoefficientsforthewingsandbodyfromthetwodifferentmeshes,whereCZand
CTarefortheverticalforceandthrustofonewing,respectively,andCZ,bandCD,bareforthevertical
forceanddragofthebody,respectively.
3.1Aerodynamicforces
Theforceandpowercoefficientsareshowninfigure8,whichincludebothinstantaneousandphase
averageddata.Thecycleaverageddataarelistedintable3foranentirecycleandfordownstroke/upstroke
separately.Figure8ashowsthattheweightsupportismostlygeneratedduringdownstrokewhereCZis
positive.Middownstrokecorrespondstothemaximumliftproduction.Duringsupinationandearlyupstroke,
Article
thewingsarestillabletogeneratesomeweightsupport.Aroundmidupstroke,verticalforcebecomes
Abstract
negativeeventhoughitsamplitudeisnotparticularlyhigh.
1.Introduction
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Figures&Data
Info&Metrics
PDF
Downloadfigure
Openinnewtab
Downloadpowerpoint
Figure8.
Forceproductionandaerodynamicpowerconsumptionofeachhummingbirdwing:(a)verticalforce
coefficient,(b)thrustcoefficientand(c)powercoefficient.Ineachcase,thethindashedlinesarethe
instantaneousdata,andthethicklineisthephaseaverageddata.
Table3. Viewinline
Forceproductionandpowerofupstroke,downstrokeandentirecycle(i.e.averagebetweenupstroke
anddownstroke).
Ontheotherhand,figure8bshowsthatthrustismostlypositiveduringbothdownstrokeandupstroke.
Furthermore,thrusthasagreaterpeakduringupstrokethanduringdownstroke.However,thedatain
table3showthatdownstrokeonaverageproducesmorethrustthanupstroke.
Figure8cshowsthatthepowerconsumptionduringbothhalfwingbeatsaresignificant.However,the
powerrequirementisgreaterfordownstroke,abouttwiceashighasupstroke.Thisfeatureissimilarto
hovering,wheredownstrokepowerisnearly2.8timesofupstrokepoweraccordingtoSongetal.[5],who
studiedtherubythroatedhummingbird.UsingtheequationP =
1
CP U
3
(2S ) fortheaerodynamic
2
Article
power,P,wehaveP=94.5mWforthecalliopehummingbird,andbodymassspecificpower34Wkg 1.Thus,
Abstract
massspecificpoweroutputofthehummingbirdiswithintherangereportedforlargerbirdspecies.For
1.Introduction 1at5ms1to47Wkg1at14ms1,anddovepower
example,cockatielpoweroutputrangesfrom17Wkg
outputrangesfrom31Wkg1at7ms1to54Wkg1at17ms1[31].Comparedwiththehoveringruby
throatedhummingbirdat55Wkg1[5],forwardflightinthecalliopehummingbirdrequireslesspower,which
4.Conclusion
isexpectedsincehoveringgenerallyismoreenergydemandingthanforwardflight.Ontheotherhand,
Ethics
forwardflightat8.3ms1isnotnecessarilyminimumpowerspeedforthehummingbird,asthemechanical
Dataaccessibility
poweroutputofbirdscanbedescribedasaUshapedcurvefunctionoftheflightspeed[3133].
Competinginterests
Funding
Usingthepresentforcecoefficientsandequation Ftotal =
1
(2CZ + CZ,b )U
2
S ,weobtainthetotal
2
Acknowledgements
verticalliftproducedbythebird,whichisaround94%ofthebirdweight.Thebirdbodyitselfgeneratesabout
References
22.2%ofbodyweight.Thisresultwillbediscussedlater.Thethrustgeneratedonthetwowingstogetheris
Leaveacomment
152%ofthebodydrag.Theimbalanceoftheverticalandhorizontalforcescouldhavebeencausedby
Figures&Data
severalreasons:(i)theabsenceofcamberinthewingmodel,(ii)errorindigitizationofthewingposition,(iii)
Info&Metrics
beakfeederinteractionasthebirdwasattemptingtofeedduringtherecording,(iv)interactionbetweenthe
PDF
birdsbodyandthewakeofthefeeder,and(v)anunderestimateofdragcoefficientforthebody.
3.2Forceproductionmechanism
Overallforceproductionofthehummingbirdcanbeexplainedfromthewingkinematicsasviewedfromthe
globalcoordinatesystem,i.e.thecoordinatesystemfixedwiththeambientair.Figure9showstheproximal
anddistalchordmovingintheglobalcoordinatesystemwiththeirtrajectoriestracedout.Duringdownstroke,
theangleofattackispositiveforboththeproximalchordandthedistalchord.Therefore,bothwingsections
generateweightsupport.Sincetheleadingedgeofthedistalsectiontiltsdownward,theaerodynamiclifthas
aforwardcomponentthatleadstothrustgenerationduringdownstroke.
Article
Abstract
1.Introduction
Downloadfigure
Openinnewtab
4.Conclusion
Ethics Downloadpowerpoint
Dataaccessibility
Figure9.
Competinginterests
Timedependentpositionofthedistalchord(a)andproximalchord(b)intheglobalcoordinatesystem
Funding
withqualitativeforceproductionatmiddownstrokeandmidupstroke.
Acknowledgements
References
Duringupstroke,bothwingsectionsmoveforwardinair,eventhoughthestrokeplaneangleislessthan90
Leaveacomment
andthewingsmovebackwardwithrespecttothebody.Nevertheless,positivethrustisgeneratedduringthis
Figures&Data
halfcyclebythedistalsection.Asshowninfigure9,theangleofattackofthedistalchordisnegativeat
Info&Metrics
upstroke,andtheoverallforceonthesectionpointsdownwardandforward.
PDF
Figure10showsthepressuredistributionwithinfourselectedverticalslicesatmiddownstrokeandmid
upstroke.Itcanbeseenthattherolesofthedistalsectionandproximalsectionaredifferent.Forboth
downstrokeandupstroke,theproximalwinghaspressuresurfaceontheventralsideandsuctionsurfaceon
thedorsalside.Thus,itsmainroleisforverticalforcegeneration.However,thedistalwingflipsitsangleof
attackbetweenthetwohalfcycles.Thus,positive(negative)pressureisdistributedontheventral(dorsal)
sideduringdownstroke,andtheoppositeistrueduringupstroke.Thispressuredifferentialleadstoweight
supportduringdownstrokeonly,butthrustproductionduringbothdownstrokeandupstroke.
Downloadfigure
Article
Abstract
Openinnewtab
1.Introduction
Downloadpowerpoint
Figure10.
4.Conclusion
Ethics
Pressuredistribution(inPascals)intheparasagittalplanesat(a)middownstrokeand(b)midupstroke.
Dataaccessibility
3.3Vortexstructures
Competinginterests
Funding
Vortexstructures,whichareinducedbythewingmotionanddominatethewake,havebeenafocalpointin
Acknowledgements
thestudyofforceproductionofflappingwingsandfishfins.Theycanalsobeusedtoevaluatewhethera
References
birdadoptsslowgaitorfastgait[22].Ashasbeenpointedoutbypreviousresearchers,atslowgaitthe
Leaveacomment
trailingedgevortices(TEVs)formringsaftereachdownstroke,andthesequencewouldlooklikeaseriesof
Figures&Data
smokerings[34,35]whileatfastgait,thetipvortices(TVs)formundulatingvortextubesfromthetip,and
Info&Metrics
theTEVsformcylindersfromthetrailingedges,bothbeingconvecteddownstream[24,25,36].
PDF
Inthecurrentstudyofhummingbirdflight,weusedtheisosurfacetoshowthevortexstructures.Thescalar
quantityisdefinedasthemaximumvalueoftheimaginarypartoftheeigenvalueofthevelocitygradient
tensoru,anditdescribesthestrengthofthelocalrotationoffluid[37].ItisfoundthattheTVsare
continuouslyshedfromthewingtip,andtheTEVsshedwiththeshapeofseparatecylinders.Suchladder
typevortexstructureswerealsoobservedinthemeasurementofforwardflightofswifts[25],whichare
closelyrelatedtohummingbirds,andlikehummingbirds,donotflextheirwingsduringflight.Aspointedout
bytheauthors[25],theladdertypewakeisformedbycontinuoussheddingofthespanwisevorticesfrom
thewingsurfaceandisdifferentfromanearlierspeculationthatforswiftsandhummingbirds,aladderlike
wakewouldbegeneratedbyadistinctvortexineachofdownstrokeandupstroke[38].Eventhoughthey
bothgeneratecontinuousladderlikevortexstructures,swiftsdonotforegoweightsupportonupstrokelike
wereportforhummingbirds.
Severalsnapshotsoftheflowfieldareshowninfigure11.Thesesnapshotsshowroughlytheshapeofthe
TVsthatfollowthetrajectoryofthewingtips.Inaddition,vortexsheddingfromthetrailingedgeisevident.
Formationoftheleadingedgevortices(LEVs)duringbothdownstrokeandupstrokeisvisible,andtheLEVs
arestableforbothdownstrokeandupstroke.ThebehaviouroftheLEVshastodowithboththe
instantaneousangleofattackandpitchingrotationofthewing[39].Fromfigure5d,theangleofattackof
thedistalsectionkeepsamaximumvaluearound25forasignificantperiodoftime,whichwouldhave
causedLEVsheddingandstall,ifthewingsimplytranslatedwithoutchangingitspitch.However,thewings
arealsoperformingrapidpitchingaroundtheiraxes,asseenfromvariationofthechordangleplottedin
figure5b.Thatis,thechordanglemagnitudedecreasesduringdownstrokeaftert/T>0.2,andquickly
increasesduringupstrokebeforemidupstroke.Suchrotationalmotionhasbeenknowntomaintainstability
ofLEVsandtoenhanceliftproductionofthewings[39].
Article
Abstract
1.Introduction
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Figures&Data
Info&Metrics
PDF
Downloadfigure
Openinnewtab
Downloadpowerpoint
Figure11.
Vortexstructuresintheflow:(a)pronation,(b)middownstroke,(c)supinationand(d)midupstroke.See
theelectronicsupplementarymaterialforananimation.
3.4Forcesonthebirdbody
Liftanddragonthebirdbodyareaffectedbytheorientationofthebirdduringflight.Ingeneral,the
inclinationangleofthebodydecreaseswiththeincreaseofflightspeed[12,23,27,40,41].Inthecurrent
study,thebodyangleofthehummingbirdisb=12,whichisclosetotheangleoftherufoushummingbirdat
aspeedof8ms1,whereb=11[12].Figure12showsboththeinstantaneousandphaseaverageddataof
theforcesonthebody.Downstroke,upstrokeandcycleaverageddataarelistedintable3.Theseresults
showthattheliftonthebodyprovides22.2%oftheweightsupport.Furthermore,liftonthebodyduring
downstrokeis1.76timesofthatduringupstroke.Infigure12a,liftonthebodyoscillatessignificantlyduring
awingbeatcycle.Ontheotherhand,dragonthebodydoesnotvaryverymuchinacycleandisnearly
equalonaveragebetweendownstrokeandupstroke,whicharereflectedinfigure12bandtable3.
Article
Abstract
1.Introduction
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Figures&Data Downloadfigure
Info&Metrics
PDF Openinnewtab
Downloadpowerpoint
Figure12.
Instantaneous(thindashedlines)andphaseaveraged(thickline)lift(a)anddrag(b)onthebirdbody.
Thinsolidlinesdenotedatafromtheisolatedbodysimulation.
Thehighpercentageofliftproducedbythebodyandtheoscillationsofthebodyliftinacyclemaybe
attributedtoaerodynamicinteractionbetweenthewingsandthebodythatisassumedtobestationaryinthe
currentstudy.Toverifythispossibility,wealsosimulatedseparatelythesameflowaroundtheisolatedbody
withoutthewingsattached.Theshapeandorientationofthebodyremainthesameinthetest.
Figure13ashowsthepressuredistributiononthebirdbodyfromtheisolatedbodysimulation,whichcanbe
comparedwiththeresultfromthefullbodysimulationshowninfigure13bandcformiddownstrokeand
midupstroke,respectively.Fortheisolatedbody,eventhoughahighpressurezoneisestablishedbelowthe
bodyandnearthehead,theflowpassesaroundthebodyintheabsenceofthewingsandmergesbehind
andabovethebody,wherethepressureispartiallyrecovered.Asaresult,theoverallliftbythebodyis
small.Whenthewingsarepresent,theflowfrombelowispreventedfrompassingaroundthebodybythe
wings.Furthermore,thewingwinginteractionmechanism,similartothatproposedbyLehmannetal.[42],
apparentlyhasplayedarolehere.Thatis,whenthetwowingsareseparatingfromoneanotherfrom
pronationtomiddownstroke,theycreatealowpressurezoneabovethebirdbody,asshowninfigure13b,
thusleadingtoanetupwardforce.Thismechanismalsoexplainswhyduringupstroke,thelowpressure
zoneabovethebody,asplottedinfigure13c,issignificantlysmallerwhencomparedwithdownstroke.Close
We use cookies to help us improve this website. Learn more
Article
Abstract
1.Introduction
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Figures&Data
Info&Metrics
PDF
Downloadfigure
Openinnewtab
Downloadpowerpoint
Figure13.
PressuredistributiononthebodysurfaceinPascals:(a)isolatedbodysimulation,(b)bodywithwingsat
middownstrokeand(c)bodywithwingsatmidupstroke.
Thepresentresultshowsthatthebirdbodyhassignificantcontributiontotheoverallweightsupport.In
comparison,previousexperimentalstudiesofinsectsandotherbirdsindicatedthatthebodyliftisonlya
smallportionoftheanimalweight,asshownintable4.However,wepointoutthatinthosepreviousstudies,
theforcewasmeasuredfortheisolatedanimalbodyonly,whileinthecurrentstudy,thewingsarepresent
andareinconstantmotion.Fortheisolatedhummingbirdbody,wealsoobservedlowliftproduction.As
shownintable4,liftoftheisolatedhummingbirdbodyisonly8%oftheweightandiscomparablewith
previousdataforinsectsandalsobirds(e.g.1520%duringflexedwingboundsinzebrafinch).
Table4. Viewinline
Liftcontributionfromthebodytoweightsupportfordifferentspecies.(Measurementfortheinsectswas
doneonisolatedbodiesandforzebrafinchwasdoneusinglivebirdswithintactbutfoldedwingsand
tails.)
3.5Comparisonofhummingbirdswithotherflyinganimals
TheadvanceratioJandstrokeplaneanglearetwoprimaryfactorsthataffecttheforceproductionof
flappingwingsduringforwardflight.Thesetwovariablesdifferlargelyamonganimalspecies.Figure14
Article
showsafewspeciesontheJmapwiththedatadirectlycollectedorderivedfromvarioussources.Itcan
Abstract
beseenthathummingbirdslargelyfallwithintherangeoftheinsectsbutalsoextendintotherangeofother
1.Introduction
birds.Forallspecies,thestrokeplaneangleincreaseswiththeadvanceratio,whichisexpectedsinceatfast
flightspeed,theanimalsnotonlyreducethebodyanglefordragreduction,whichwouldnaturallycausethe
strokeplaneangletoincrease,butmayalsotiltthestrokeplanefurthertoenhancethrustgeneration.
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Figures&Data
Info&Metrics
PDF
Downloadfigure
Openinnewtab
Downloadpowerpoint
Figure14.
(a)Strokeplaneangleversusadvanceratio.Dataarecollectedfromtheliterature(Diptera:[48],
bumblebee:[15,17],rufoushummingbird:[12],magpie,pigeonandzebrafinch:[40,41],hawkmoth:[49],
barnswallow:[50],bat:[51]).
Forthesmallinsectslikebumblebeesandfruitflies,theadvanceratioisusuallylessthanone[14].Forsuch
slowflight,liftproductionispredominantoverthrustproduction.Sincethebacksweepingvelocityofthe
distalwingexceedstheforwardflightspeedatupstroke[15,20,33],thewingtiptrajectorytracedoutinthe
globalcoordinatesystemishighlybackwardskewedatupstroke,whichisshowninfigure15fora
bumblebeeatJ=0.6.Inthiscase,downstrokeismainlyforliftproduction,whileupstrokeismainlyforthrust
production.Iftheflightspeedisfurtherreduced,withamoreskewedtrajectory,upstrokemayevenproduce
liftaswell.Overall,thisstrategyofusingupstrokeisalsoknownasbackwardflick[18,23,31].Anexception
isthefruitflywhichisshownbyarecentstudythatitsupstrokeusesapaddlingmodetoproducedrag
Article
basedthrust[20].
Abstract
1.Introduction
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Figures&Data
Info&Metrics
PDF
Downloadfigure
Openinnewtab
Downloadpowerpoint
Figure15.
Wingtiptrajectoryandforceproductionofbumblebee,hummingbirdandpigeon.
Formostbirds,theadvanceratioissignificantlygreaterthanone,andthestrokeplaneangleiscloseto90.
Thus,thewingtiptrajectoryintheglobalcoordinatesystembecomesmuchlessbackwardskewedandthe
wavelengthtoamplituderatiobecomesgreaterasshowninfigure15forapigeon.Inthiscase,upstrokeis
notsuitableforthrustgeneration.Instead,thewingsareeitherfeatheredwithlittleforceproduced[25,24],or
sweptonupstrokewithsomeliftproduction(e.g.pigeon)[26,27].Ontheotherhand,apowerfuldownstroke
isusedtoproducebothliftandthrust.
Forthehummingbirdinthecurrentstudy,theadvanceratioisbetweenthatofinsectsandotherbirds.The
wingtiptrajectoryismoderatelyskewedasshowninfigure15.Therefore,withproperangleofattack,the
wingscanstillproducethrustduringupstroke.However,sincetheoverallforcepointsdownward,somelift
hastobesacrificed.Fromthisfigure,itcanbeseenthatthrustcanbeproducedwhenthewingspeedat
upstrokeiscomparabletoorpossiblyevenlowerthantheflightspeed.Thisthrustmechanismisanalogous
toasailthatmovesagainstwindandthusistermedasailmodeinthiswork.Ontheotherhand,
downstrokeofthehummingbirdissimilartothatofbigbirds,asshowninfigure15,wherebothliftand
Article
thrustaregenerated.
Abstract
1.Introduction
Itshouldbepointedoutthatatslowflightspeeds,forceproductionofthehummingbirdstillappearscloseto
thatofinsects.AsshownbyTobalskeetal.[12],whenJisbelow0.7,thestrokeplaneangleofthe
hummingbirdissmallandthewingtiptrajectoryisalsohighlyskewedlikethatofinsects.Similarly,some
4.Conclusion
insectscanalsoperformfastflightatJ>1,e.g.hawkmoth,asseenfromfigure14.Itwouldbeinterestingto
Ethics
seewhethertheirforceproductionmechanismissimilartothatdescribedhereforthehummingbird.
Dataaccessibility
Competinginterests
Thereissignificantsimilaritybetweenthehummingbirdandbats,bothbeingcapableofhoveringand
Funding
forwardflightatJ=1[52].Recentflowvisualizationstudiesofbatshavesuggestedthataerodynamicfunction
Acknowledgements
ofupstrokechangeswithforwardflightspeed[16].Athoveringandslowspeeds,batwingsareinverted
References
duringupstroketoaidweightsupportandatfastspeeds,downstrokeproducesweightsupportandthrust,
Leaveacomment
butupstrokemaygenerateextrathrustatcostofnegativelift.Thesefeaturesaresimilartowhatwehave
Figures&Data
reportedforthehummingbird,whichisinterestinggiventhat,unlikehummingbirds,batsflextheirwings
Info&Metrics
duringupstroke.
PDF
Itisalsointerestingtopointoutthesimilaritiesanddifferencesbetweenthehummingbirdandtheswift,as
bothofthemkeeptheirwingsextendedduringtheentirewingbeatcycleregardlessofflightspeeds.Thisis
highlyunusualforbirds,asmostbirdsflexthewingsduringupstroke[53].Regardlessofkeepingthewings
extended,thecombinationofspanwisetwistandthestrokeplaneanglereportedhereforthehummingbird
isamechanismforpermittingflightoverawiderangeofspeed.Fortheswift,itisbelievedthatwingtwistis
animportantfactorleadingtooptimalefficiencyofflappingflight[54].Ontheotherhand,theswiftsflight
styleisdedicatedtocruisingflight,anditisnotadeptathovering,andsomeswiftspeciesareevenunableto
accelerateintoflightfromastandingstartwithoutfirstdroppinginaltitudetogainvelocity[55].Therefore,
oneconclusioncouldbethattheswiftpatternofforcedevelopmentisoptimizedformorecontinuousweight
supportduringcruisingflightinawaythatthecloselyrelatedhummingbirddoesnotachievewhileforthe
hummingbird,itsevolutionarytrajectorymayhavefavouredthecapacityforsomethrustproductionatthe
expenseofthemoreconstantweightsupportstyleintheswift[54].Thishypothesismeritstestingina
broaderphylogeneticcontext,asitisnotappropriatetoinferevolutionarytrajectoriesfromtwospecies
comparisons[56].
4.Conclusion
Threedimensionalcomputersimulationhasbeenperformedtostudyaerodynamicsofahummingbirdinfast
forwardflightandwhosewingmotionwascapturedbyfilmingthebirdflightinawindtunnel.Thefinding
placeshummingbirdsinaninterestingpositionrelativetoinsectsandlargebirds.Ataspeedof8.3ms1,the
advanceratioofthehummingbirdisbetweenthoseoftypicalinsectsandlargebirds,andthesimulation
resultsshowthatthehummingbirdusesadifferentstrategyforliftandthrustproduction.Inparticular,its
We use cookies to help us improve this website. Learn more
powerdownstrokegeneratesbothweightsupportandthrustjustlikeotherbirds,butitsupstrokefurther Close
enhancesthrustbysettingthedistalwingsataproperangleofattackwithrespecttotheoncomingair,even
thoughsuchthrustenhancementcomesatcostofsomenegativeliftatupstroke.Thesefeaturesarethus
Article
similartothoseofbatsflyingatJ=1.Ultimately,cautionisnecessaryininterpretingourresultsasthey
Abstract
emanatefromthestudyofasinglebirdofonespecies.Newstudiesinaphylogeneticcontextwillbeuseful
1.Introduction
forunderstandingtheevolutionarytrajectoriesandselectivepressuresthatdrovethehummingbirdflight
style. 3.Resultsanddiscussion
4.Conclusion
Ethics
Ethics
Dataaccessibility
Competinginterests
AllprotocolsassociatedwithhummingbirdcareandexperimentationwereapprovedbytheUniversityof
Funding
MontanaIACUC(06314BTDBS12914).ScientificcollectingwaspermittedbytheUSFishandWildlife
Acknowledgements
Service(SCCL771277)andMontanaDepartmentofFishWildlifeandParks(2015062).
References
Leaveacomment
Dataaccessibility
Figures&Data
Info&Metrics
DataavailablefromtheDryadDigitalRepository:http://dx.doi.org/10.5061/dryad.8ch1b.
PDF
Competinginterests
Wedeclarewehavenocompetinginterests.
Funding
ThisresearchwassupportedbytheNSF(no.CBET0954381toH.L.andno.CMMI1234737toB.T.).
Acknowledgements
ThecomputingresourceswereprovidedbytheNSFXSEDEandtheVanderbiltACCRE.
ReceivedMarch31,2016.
AcceptedMay10,2016.
2016TheAuthors.PublishedbytheRoyalSocietyunderthetermsoftheCreativeCommonsAttribution
Licensehttp://creativecommons.org/licenses/by/4.0/,whichpermitsunrestricteduse,providedtheoriginal
authorandsourcearecredited.
References
1.WarrickDR,TobalskeBW,PowersDR.2005Aerodynamicsofthehoveringhummingbird.Nature 435,10941097.
We useCrossRef
(doi:10.1038/nature03647) cookiesMedline
to helpWebofScience
us improve this website. Learn more Close
2.WarrickDR,TobalskeBW,PowersDR.2009Liftproductioninthehoveringhummingbird.Proc.R.Soc.B276,37473752.
Article
(doi:10.1098/rspb.2009.1003) Abstract/FREEFullText
Abstract
3.WolfM,OrtegaJimenezVM,DudleyR.2013StructureofthevortexwakeinhoveringAnnashummingbirds(CalypteAnna).
1.Introduction
Proc.R.Soc.B280,20132391.(doi:10.1098/rspb.2013.2391) Abstract/FREEFullText
4.KruytJW,QuicaznRubioEM,vanHeijstGF,AltshulerDL,LentinkD.2014Hummingbirdwingefficacydependsonaspect
4.Conclusion
ratioandcompareswithhelicopterrotors.J.R.Soc.Interface 11,20140585.(doi:10.1098/rsif.2014.0585)
Abstract/FREEFullText
Ethics
Dataaccessibility
5.SongJ,LuoH,HedrickTL.2014Threedimensionalflowandliftcharacteristicsofahoveringrubythroatedhummingbird.J.
Competinginterests
R.Soc.Interface 11,20140541.(doi:10.1098/rsif.2014.0541) Abstract/FREEFullText
Funding
Acknowledgements
6.SongJ,LuoH,HedrickTL.2015Wingpitchingmechanismofhoveringrubythroatedhummingbirds.Bioinspir.Biomimet.
10,016007.(doi:10.1088/17483190/10/1/016007)
References
CrossRef
Leaveacomment
7.EllingtonCP,vandenBergC,WillmottAP,ThomasALR.1996Leadingedgevorticesininsectflight.Nature 384,626630.
Figures&Data
(doi:10.1038/384626a0) CrossRef WebofScience
Info&Metrics
8.FrySN,SayamanR,DickinsonMH.2005TheaerodynamicsofhoveringflightinDrosophila.J.Exp.Biol.208,23032318.
PDF
(doi:10.1242/jeb.01612) Abstract/FREEFullText
9.ZhengL,HedrickTL,MittalR.2013Amultifidelitymodellingapproachforevaluationandoptimizationofwingstroke
aerodynamicsinflappingflight.J.FluidMech.721,118154.(doi:10.1017/jfm.2013.46) CrossRef WebofScience
10.AonoH,LiangF,LiuH.2008Nearandfarfieldaerodynamicsininsecthoveringflight:anintegratedcomputationalstudy.
J.Exp.Biol.211,239257.(doi:10.1242/jeb.008649) Abstract/FREEFullText
11.HedrickTL,TobalskeBW,RosIG,WarrickDR,BiewenerAA.2012Morphologicalandkinematicbasisofthehummingbird
flightstroke:scalingofflightmuscletransmissionratio.J.R.Soc.Interface 279,19861992.(doi:10.1098/rspb.2011.2238)
12.TobalskeBW,WarrickDR,ClarkCJ,PowersDR,HedrickTL,HyderGA,BiewenerAA.2007Threedimensionalkinematics
ofhummingbirdflight.J.Exp.Biol.210,23682382.(doi:10.1242/jeb.005686) Abstract/FREEFullText
13.VogelS.1966FlightinDrosophila.I.Flightperformanceoftetheredflies.J.Exp.Biol.44,567578. Abstract/FREEFullText
14.DudleyR.2002Thebiomechanicsofinsectflight:form,function,evolution .Princeton,NJ:PrincetonUniversityPress.
15.DudleyR,EllingtonCP.1990Mechanicsofforwardflightinbumblebee.I.Kinematicsandmorphology.J.Exp.Biol.148,
1952. Abstract/FREEFullText
16.HedenstrmA,JohanssonLC.2015Batflight:aerodynamics,kinematicsandflightmorphology.J.Exp.Biol.218,653663.
(doi:10.1242/jeb.031203) Abstract/FREEFullText
17.DudleyR,EllingtonCP.1990Mechanicsofforwardflightinbumblebees.II.Quasisteadyliftandpowerrequirements.J.
Exp.Biol.148,5388. Abstract/FREEFullText

18.NorbergUM.1976Aerodynamics,kinematics,andenergeticsofhorizontalflappingflightinthelongearedbatPlecotus
auritus.J.Exp.Biol.65,179212. Abstract/FREEFullText
Article
19.CrandellKE,TobalskeBW.2015Kinematicsandaerodynamicsofavianupstrokesduringslowflight.J.Exp.Biol.218,
Abstract
25182527.(doi:10.1242/jeb.116228) Abstract/FREEFullText
1.Introduction
20.RistrophL,BergouAJ,GuckenheimerJ,WangZJ,CohenI.2011Paddlingmodeofforwardflightininsects.Phys.Rev.Lett.
106,178103.(doi:10.1103/PhysRevLett.106.178103) CrossRef Medline
4.Conclusion
21.PennycuickCJ.1990Predictingwingbeatfrequencyandwavelengthofbirds.J.Exp.Biol.150,171185.
Ethics
Dataaccessibility
22.AlexanderDE,VogelS.2004Naturesflyers:birds,insects,andthebiomechanicsofflight.Baltimore,MA:JHUPress.
Competinginterests
Funding
23.TobalskeBW,HedrickTL,BiewenerAA.2003Wingkinematicsofavianflightacrossspeeds.J.AvianBiol.34,177184.
Acknowledgements
(doi:10.1034/j.1600048X.2003.03006.x) CrossRef WebofScience
References
24.SpeddingGR,RosnM,HedenstrmA.2003Afamilyofvortexwakesgeneratedbyathrushnightingaleinfreeflightina
Leaveacomment
windtunneloveritsentirenaturalrangeofflightspeeds.J.Exp.Biol.206,23132344.(doi:10.1242/jeb.00423)
Figures&Data
Info&Metrics
25.HenningssonP,SpeddingGR,HedenstrmA.2008Vortexwakeandflightkinematicsofaswiftincruisingflightinawind
PDF
tunnel.J.Exp.Biol.211,717730.(doi:10.1242/jeb.012146) Abstract/FREEFullText
26.HedrickTL,TobalskeBW,BiewenerAA.2002Estimatesofcirculationandgaitchangebasedonathreedimensional
kinematicanalysisofflightincockatiels(Nymphicushollandicus)andringedturtledoves(Streptopeliarisoria).J.Exp.Biol.
205,13891409. Abstract/FREEFullText
27.HedrickTL,UsherwoodJR,BiewenerAA.2004Winginertiaandwholebodyacceleration:ananalysisofinstantaneous
aerodynamicforceproductionincockatiels(Nymphicushollandicus)flyingacrossarangeofspeeds.J.Exp.Biol.207,
16891702.(doi:10.1242/jeb.00933) Abstract/FREEFullText
28.TobalskeBW,PuccinelliLA,SheridanDC.2005Contractileactivityofthepectoralisinthezebrafinchaccordingtomode
andvelocityofflapboundingflight.J.Exp.Biol.208,28952901.(doi:10.1242/jeb.01734) Abstract/FREEFullText
29.HedrickTL.2008Softwaretechniquesfortwoandthreedimensionalkinematicmeasurementsofbiologicaland
biomimeticsystems.Bioinspir.Biomimet.3,034001.(doi:10.1088/17483182/3/3/034001) CrossRef
30.LuoH,DaiH,FerreiradeSousaP,YinB.2012Onnumericaloscillationofthedirectforcingimmersedboundarymethod
formovingboundaries.Comput.Fluids56,6176.(doi:10.1016/j.compfluid.2011.11.015) CrossRef WebofScience
31.TobalskeBW,HedrickTL,DialKP,BiewenerAA.2003Comparativepowercurvesinbirdflight.Nature 421,363366.
(doi:10.1038/nature01284) CrossRef Medline
32.PennycuickCJ.1968PowerrequirementsforhorizontalflightinthepigeonColumbalivia.J.Exp.Biol.49,527555.
(doi:10.1098/rsif.2014.0953) Abstract/FREEFullText
33.SunM,WuJH.2003Aerodynamicforcegenerationandpowerrequirementsinforwardflightinafruitflywithmodeledwing
motion.J.Exp.Biol.206,30653083.(doi:10.1242/jeb.00517) Abstract/FREEFullText
34.SpeddingGR,RaynerJMV,PennycuickCJ.1984Momentumandenergyinthewakeofapigeon(Columbalivia)inslow
We use cookies
flight.J.Exp.Biol.111,81102. to help us improve this website. Learn more
Close
35.SpeddingGR.1986Thewakeofajackdaw(Corvusmonedula)inslowflight.J.Exp.Biol.125,287307.
Article
Abstract
36.SpeddingGR.1987Thewakeofakestrel(Falcotinnunculus)inflappingflight.J.Exp.Biol.127,5978.
1.Introduction
37.MittalR,BalachandarS.1995Generationofstreamwisevorticalstructuresinbluffbodywakes.Phys.Rev.Lett.75,1300.
4.Conclusion
(doi:10.1103/PhysRevLett.75.1300) CrossRef Medline WebofScience
Ethics
38.PennycuickCJ.1988Onthereconstructionofpterosaursandtheirmannerofflight,withnotesonvortexwakes.Biol.Rev.
Dataaccessibility
63,299331.(doi:10.1111/j.1469185X.1988.tb00633.x) CrossRef GeoRef
Competinginterests
Funding
39.EldredgeJD,ToomeyJ,MedinaA.2010Ontherolesofchordwiseflexibilityinaflappingwingwithhoveringkinematics.J.
Acknowledgements
FluidMech.659,94115.(doi:10.1017/S0022112010002363) CrossRef WebofScience
References
40.TobalskeB,DialK.1996Flightkinematicsofblackbilledmagpiesandpigeonsoverawiderangeofspeeds.J.Exp.Biol.
Leaveacomment
199,263280. Abstract/FREEFullText
Figures&Data
Info&Metrics
41.TobalskeBW,PeacockWL,DialKP.1999Kinematicsofflapboundingflightinthezebrafinchoverawiderangeofspeeds.
J.Exp.Biol.202,17251739. Abstract
PDF
42.LehmannFO,SaneSP,DickinsonM.2005Theaerodynamiceffectsofwingwinginteractioninflappinginsectwings.J.
Exp.Biol.208,30753092.(doi:10.1242/jeb.01744) Abstract/FREEFullText
43.NachtigallW.1964Zuraerodynamikdescoleopterenfluges:wirkendieelytrenalstragflchen.Verhandlungender
DeutschenZoologischenGesellschaft27,319326.
44.WoodJ.1970AstudyoftheinstantaneousairvelocitiesinaplanebehindthewingsofcertainDipteraflyinginawindtunnel.
J.Exp.Biol.52,1725. Abstract/FREEFullText
45.ChanceMAC.1975Airflowandtheflightofanoctuidmoth.InSwimmingandflyinginnature(edsTYTWu,CJBrokaw,C
Brennen),pp.829843.Berlin,Germany:Springer.
46.CsicskyMJ.1977Bodyglidinginzebrafinch.FortschrittederZoologie 24,275286. WebofScience
47.TobalskeBW,HearnJWD,WarrickDR.2010Aerodynamicsofintermittentboundsinflyingbirds.Exp.Fluids46,963973.
(doi:10.1007/s0034800906149)
48.EnnosAR.1989ThekinematicsandaerodynamicsofthefreeflightofsomeDiptera.J.Exp.Biol.142,4985.
49.WillmottAP,EllingtonCP.1997ThemechanicsofflightinthehawkmothManducasexta.I.kinematicsofhoveringand
forwardflight.J.Exp.Biol.200,27052722. Abstract/FREEFullText
50.ParkKJ,RosnM,HedenstrmA.2001Flightkinematicsofthebarnswallow(Hirundorustica)overawiderangeofspeeds
inawindtunnel.J.Exp.Biol.204,27412750. Abstract/FREEFullText
51.WolfM,JohanssonLC,vonBusseR,WinterY,HedenstrmA.2010Kinematicsofflightandtherelationshiptothevortex
wakeofaPallaslongtonguedbat(Glossophagasoricina).J.Exp.Biol.213,21422153.(doi:10.1242/jeb.029777)
We
Abstract/FREEFullText use cookies to help us improve this website. Learn more Close
52.HedenstrmA,MuijresFT,BusseR,JohanssonLC,WinterY,SpeddingGR.2009HighspeedstereoDPIVmeasurement
Article
ofwakesoftwobatspeciesflyingfreelyinawindtunnel.Exp.Fluids46,923932.(doi:10.1007/s0034800906345)
WebofScience
CrossRef Abstract
1.Introduction
53.HenningssonP,BomphreyRJ.2011Timevaryingspanefficiencythroughthewingbeatofdesertlocusts.J.R.Soc.
Interface 9,20110749.(doi:10.1098/rsif.2011.0749)
4.Conclusion
54.HenningssonP,HedenstrmA,BomphreyRJ.2014Efficiencyofliftproductioninflappingandglidingflightofswifts.PLoS
ONE9,e90170.(doi:10.1371/journal.pone.0090170)
Ethics CrossRef Medline
Dataaccessibility
55.WarrickDR.1998Theturningandlinearmaneuveringperformanceofbirds:thecostofefficiencyforcoursinginsectivores.
Competinginterests
Can.J.Zool.76,10631079.(doi:10.1139/z98044) CrossRef
Funding
Acknowledgements
56.GarlandT,AdolphSC.1994Whynottodotwospeciescomparativestudies:limitationsoninferringadaptation.Physiol.
Zool.67,797828.(doi:10.1086/physzool.67.4.30163866)
References CrossRef
Leaveacomment
ViewAbstract
Figures&Data
Info&Metrics
PDF
0Comments RoyalSocietyOpenScience
1 Login
Recommend Share SortbyNewest
Startthediscussion
Bethefirsttocomment.
ALSOONROYALSOCIETYOPENSCIENCE
Rapidmimicryandemotionalcontagionin Beautyisintheefficientcodingofthe
domesticdogs beholder
Previous Next
Backtotop
Previous
Article Next
Abstract
June2016
1.Introduction
TableofContents
AbouttheCover
Indexbyauthor
4.Conclusion
Ethics
Dataaccessibility
Competinginterests
Funding
Acknowledgements
References
Leaveacomment
Searchthisissue
Figures&Data
Info&Metrics
KEYWORDS
PDF
flappingwings,hummingbirds,forwardflight,unsteadyaerodynamics
Share Citationtools
Email Download
Print Articlereuse
Managealerts
Seerelatedsubjectareas:
biomechanics ,fluidmechanics ,mechanicalengineering
Relatedarticles
Norelatedarticlesfound
WebofScience GoogleScholar
Citedby
LearnaboutdisplayedequationsinRoyalSocietyOpenScience
RECOMMENDEDARTICLES
Threedimensionalflowandliftcharacteristicsofahoveringrubythroatedhummingbird.
JialeiSongetal.,Interface,2014
Heatdissipationduringhoveringandforwardflightinhummingbirds
Article
KeelyR.Corderetal.,RoyalSocOpenSci,2015
Abstract
Vortexwake,downwashdistribution,aerodynamicperformanceandwingbeatkinematicsinslowflyingpiedflycatchers.
FlorianTMuijresetal.,Interface,2011
1.Introduction
Computationalinvestigationofcicadaaerodynamicsinforwardflight.
HuiWanetal.,Interface,2015
4.Conclusion
BiologyandPhysicsofLocustFlight.III.TheAerodynamicsofLocustFlight
MartinJensen,PhilosophicalTransactionsB,1956
Ethics
Dataaccessibility
Hummingbirdwingefficacydependsonaspectratioandcompareswithhelicopterrotors.
JanWKruytetal.,Interface,2014
Competinginterests
Funding
Flowvisualizationandunsteadyaerodynamicsintheflightofthehawkmoth,Manducasexta
AdrianL.R.Thomasetal.,PhilosophicalTransactionsB,1997
Acknowledgements
References
Upstrokewingflexionandtheinertialcostofbatflight.
DanielKRiskinetal.,ProceedingsB,2012
Leaveacomment
Figures&Data
Info&Metrics Poweredby
PDF
ROYALSOCIETYOPENSCIENCE
Aboutthisjournal
Contactinformation
Submit
Authorbenefits
Openpeerreview
Commentingguidelines
Datapolicy
Openaccessmembership
Recommendtoyourlibrary
FAQs
Help
ROYALSOCIETYPUBLISHING
Ourjournals
Openaccess
Publishingpolicies
Conferences
Podcasts
News
Blog
Manageyouraccount
Terms&conditions
Cookies
THEROYALSOCIETY
Article
Aboutus Abstract
Contactus
1.Introduction
Fellows
Events
Grants,schemes&awards
4.Conclusion
Topics&policy
Collections Ethics
Venuehire Dataaccessibility
Competinginterests
Funding
Acknowledgements
OnlineISSN20545703|Copyright2016TheRoyalSociety
References
Leaveacomment
Figures&Data
Info&Metrics
PDF

Three-Dimensional Simulation For Fast Forward Flight of A Calliope Hummingbird - Open Science

Transféré par

Informations du document

Titre original

Copyright

Formats disponibles

Partager ce document

Partager ou intégrer le document

Options de partage

Avez-vous trouvé ce document utile ?

Ce contenu est-il inapproprié ?

Droits d'auteur :

Formats disponibles

Three-Dimensional Simulation For Fast Forward Flight of A Calliope Hummingbird - Open Science

Transféré par

Droits d'auteur :

Formats disponibles

19/09/2016 Threedimensionalsimulationforfastforwardflightofacalliopehummingbird|OpenScience

Home Content Informationfor Aboutus Signup Submit

Article Figures&Data Info&Metrics PDF

We use cookies to help us improve this website. Learn more Close

We use cookies to help us improve this website. Learn more Close

We use cookies to help us improve this website. Learn more Close

We use cookies to help us improve this website. Learn more Close

wheref isthestressonthewingsurface,andu isthevelocityofapointonthewinginthebodyfixed

We use cookies to help us improve this website. Learn more Close

We use cookies to help us improve this website. Learn more Close

We use cookies to help us improve this website. Learn more Close

We use cookies to help us improve this website. Learn more Close

46.CsicskyMJ.1977Bodyglidinginzebrafinch.FortschrittederZoologie 24,275286. WebofScience

Recommend Share SortbyNewest

biomechanics ,fluidmechanics ,mechanicalengineering

Vous aimerez peut-être aussi