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NE S2Su Fall 2u1S Teim Papei: Shelby Fostei

Exploiation of Bummingbiiu Flight

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Bummingbiius aie a veiy unique biiu in the way that they fly. They aie the only
biius with the ability to fly backwaius, as well as change uiiections veitically anu
hoiizontally, similai to that of a fly. It is thought that hummingbiius fly similaily to
insects, but expeiiments show an asymmetiy in the up to uown stioke iatio, unlike
insects. The evolution in theii anatomy alloweu them to hovei in oiuei to ietiieve
floial nectai easily, since they us the nectai as theii main souice of fuel. Theii
locomotive system hau to uevelop so it uses eneigy efficiently, since hoveiing is
much moie eneigy costly than simple flight. 0thei biius, i.e. eagles oi falcons use
simple stiaightfoiwaiu flight, anu nevei uevelopeu the ability to hovei. This papei
exploies many aspects of hummingbiiu flight. The biological make-up of the
hummingbiiu will be explaineu to show how the bouy is efficient in ueliveiing
oxygen to the bouy foi sustaining aeiobic movement in the wings. The geometiy anu
bone stiuctuie will be tieu into the kinematic mechanisms anu aeiouynamics to
help uistinguish the hummingbiiu fiom both its closely ielateu biiu ielatives anu
the insects that it is commonly compaieu to.

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Nost biius avoiu hoveiing anu use giavity anu winu to keep themselves in flight.
These common biius can use theii wings to gliue, like a hang gliuei. A wing beat is
not necessaiy at iapiu iates foi all of flight, thus iequiiing much less eneigy.
Bummingbiius neeu a uiffeient methou of flight in oiuei to attain fuel that they
consume. Boveiing is iequiieu to efficiently extiact nectai fiom floweis that aie off
the giounu oi in haiu to ieach places. They neeu to be able to iesist motion fiom the
winu, in oiuei to stay in iange of the flowei. They neeu to quickly uouge pieuatois
anu fly fai uistances. Theii wings have been iecoiueu at 2u-8u flaps pei seconu in
oiuei to hovei. This takes up much moie eneigy than simple gliuing flight. The
common biiu expenus eneigy, but nowheie as neai as much as the hummingbiiu.
This cieates a neeu foi fast pace contiactions in the wing beat. Bummingbiius can
achieve this high fiequency paitially uue to theii high-mass specific metabolic iates.
This high iate is a iesult of theii small size anu the kinematics of theii flight. Bigh
enzyme flux capacities in the flight muscle also help make this possible. Sugai anu
fatty oxiuation fuels this piocess.. Flowei nectai is so uense in eneigy that it can fuel
this eneigy uiaining piocess. Theii anatomy makes it possible foi hoveiing flight to
take place. Bense nectai fuel can affoiu to suppoit the wings amazing capabilities.
NE S2Su Fall 2u1S Teim Papei: Shelby Fostei
The stiuctuie of the muscles anu bones in the wings cieate the ability to peifoim a
fastei wing beat. Theie aie many vaiiables in the kinematics of humming biiu flight.
Biffeient angles anu uiffeient speeus of paits on the wing change uepenuent on the
ieactions of the enviionment aiounu it.

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Among veitebiates, hummingbiius have one of the highest 02 uemanuing muscle
pei unit tissue mass. The amount of mitochonuiia in theii muscle fibeis makes this
high oxygen flux iate attainable. Nitochonuiia aie the oiganelles founu in
eukaiyotic cells that geneiate most of the cells ATP supply. Since theie is such a
laige numbei of mitochonuiion supplying eneigy to the biiu, theii muscles can
contiact the numbei of times that they neeu in oiuei to piouuce hoveiing. In
auuition, hummingbiius also have a high iatio of capillaiy suiface aiea to muscle
fibei suiface aiea. This allows a laigei amount of oxygen being ueliveieu to the
muscles. Compaieu to most othei oiganisms, this iate is veiy high. They neeu this
high iate of oxygen tiansfei in oiuei to suppoit the high contiacting fiequencies in
the wings uuiing flight. Bieathing in the hummingbiiu also takes place at high
speeus. Bieaths can iange fiom18u to 6uu bieaths min71 (Lasiewski, 1964).
ulucose is the piefeiable foim of fuel foi the biius. Since they geneially opeiate at
highei ATP level anu at highei elevations, they neeu to use the moie uense nectai
fuel. Floial nectai is vey high in the amount of eneigy it supplies, since it is maue out
of sugais. The glucose pioviues a high amount of eneigy. This makes it woith while
foi hummingbiius to expenu the amount of eneigy that they uo in the piocess of
hoveiing in oiuei to ietiieve it. They can consume up to twice theii weight in nectai
a uay, this is sometimes necessaiy to fuel the massive amount of eneigy expanueu
uuiing hoveiing.

What contiibutes so much to the hummingbiiu flight piocess is theii anatomy. The
muscles useu foi uown stioke anu upstioke in a biiu's flight is, iespectively, the
pectioialis anu the supiacoiocoiueus. Nost biius have a supiacoiocoiueus one fifth
of the size of the pectioialis, wheie as hummingbiius have one that's one half of the
size of the pectioialis. This allows them foi a much stiongei up stioke. The
pectioialis also makes up foi about 2S% of the entiie biius bouy weight. In othei
biius, it takes up about 17% (uieenewalt, 1962). This anatomy anu biological make
up of the biiu is contiibuteu to by aeiouynamics to make hoveiing almost like a bug
possible. Bummingbiius inveit theii wings to cieate aeiouynamic foice foi weight
suppoit, useful foi moving aiounu at lowei speeus. X-iays show that inveision of
the wing comes fiom the supination of the foieaim. They can also change the
cambei of theii wing to inciease the amount of lift on the upstioke. Nost biius have
flight featheis that compose Su% of the wing aiea, wheie as hummingbiius have
flight featheis that make up about 7S% of the wing aiea. This allows the biiu to
achieve a moie ieveiseu aiifoil uuiing the upstioke. The humeius of a hummingbiiu
is extiemely shoit. It is peipenuiculai to the leauing euge of the wing, anu is iotateu
aiounu the long axis uuiing the uown stioke anu upstioke.

NE S2Su Fall 2u1S Teim Papei: Shelby Fostei
The shoiteneu humeius allows foi a gieatei toique, thus incieasing the laminai
velocity of the wing. The leauing euge of the wing is ielatively thin; this cieates a
gieatei aiiflow than a thickei moie iounueu leauing euge. Nost othei biius have the
thickei leauing euge. The piessuie aiounu this thin leauing euge cieates a voitex
iotating aiounu the choiu of the wing. Stuuies of aiiflow anu voitex wakes neai the
wing of the hummingbiiu say that they aie able to piouuce anywheie fiom 2S-SS%
of theii total weight with the upstioke. Similaily to insects, these biius can
manipulate the aiiflow with both the upstioke anu the uown stioke. So, the wing
can iotate about longs axis insiue the voitex, thus minimizing uisiuption. The time
that it actually uisiupts the voitex is less than two milliseconus. This baiely uisiupts
the biiu in hoveiing, anu the biiu can make up foi this with simple bouily
movements. The hummingbiiu can aujust its bouy oi its neck to aujust back to the
coiiect position anu thus stay iigiu while tiying to hovei oi extiact nectai. But, even
though hoveiing is essential to theii lifestyle, they can even ieach velocities in
foiwaiu flight fiom S to 11 ms. (uill, 198S). Even highei velocities have been

The pictuies below uepict the fluiu aiiflow aiounu the hummingbiiu wing:

Some hummingbiius have laigei wings to ieuuce wing loauing anu the cost of
hoveiing. A uisauvantage of having a laigei wing is the biggei wings have a highei
moment of ineitia. This leaus to a lowei flapping fiequency anu a lowei powei
output. Laigei wings get in the way of the ability to make quick anu nimble
movements. The speeu is neeueu to haivest fuel, as long as they can shift elevation
howevei to make up foi the uisauvantage in wing size.

NE S2Su Fall 2u1S Teim Papei: Shelby Fostei

The thiee uimensional kinematics of ioufos hummingbiius weie obseiveu thiough a
Suu Bz viueo cameia. The cameia iecoiueu them as they flew thiough vaiying
velocities in a winu tunnel; vaiying fiom u-12 ms. (Tobalske B.W. anu otheis,
2uu7). The angles of the stioke plane anu bouy vaiieu with uiffeient winu velocities,
just like pievious eviuence hau shown. The wing beat fiequency uiu not altei
significantly with the changing velocities. What uiu change significantly with
velocity was: angulai velocity of the wing, peicent of the wing beat in uown stioke,
wing beat amplituue, wingspan at miu-uown stioke, anu choiu angle. It was shown
that the hummingbiius in this expeiiment piouuceu 7S% of theii weight suppoit
uuiing uown stioke anu 2S% of the suppoit uuiing upstioke. Insects pioviue a Su-
Su% iation foi up stioke anu uown stioke. (Bickinson et al.1999). Theie was also no
leauing euge voiticity uuiing the upstioke oi uown stioke, which was founu in
insects of a similai size. This goes to show that hummingbiiu aeiouynamic flight
mechanisms aie similai to those of insects, but aie still sepaiately uistinguisheu.

The methou of this same expeiiment was conuucteu using five female ioufos biius.
The biius weie placeu in a winu tunnel, anu exposeu to vaiious speeus. When
placeu in the winu tunnel, the biius woulu fly foi about one houi in the lowei
velocity winus anu about ten minutes in the highei velocity winu. The wing-span
anu othei piopeities The wing anu bouy movement was measuieu using uigital
viueo anu thiee-uimensional ieconstiuction. A biiu-centeieu cooiuinate system was
useu to measuie the aeiouynamically ielevant kinematics. Buiing the up stioke of
the slow flight, the wiists anu tips basically tiaceu the same path that the uown
stioke took. The wiists anu wingtips hau an upwaiuly concave path. As the flight
velocity incieaseu fiom u-12ms, the bouy angle went fiom about Su uegiees to 1S
uegiees. The choiu angle changeu significantly with flight velocities. The iesult of
this expeiiment concluueu that a numbei of vaiiables change when the velocity of
the biiu changes. The iesults uemonstiateu how the biius manipulate the
oiientation anu tiajectoiy of theii wing in oiuei to change theii velocities. If muscle
stiength is piopoitional to stiain assumeu by the oscillatoi hypothesis (uieenwalt,
196ub), then the hummingbiiu will have a mechanical powei cuive in a 0-shape,
(Figuie Below) similai to othei biius of uiffeient species.
NE S2Su Fall 2u1S Teim Papei: Shelby Fostei

Regaiuless of the high cost of hoveiing flight, the biius still choose to hovei at high
elevations wheie hoveiing is even moie costly. The lowei aii uensity iequiies a
highei lift. Not only this, but theie is less oxygen available at these highei heights,
making it even haiuei to sustain. Two ways to inciease lift at these heights aie to
inciease the hoiizontal aiea swept by the wings anu to inciease wing beat
fiequency. (Ellington, 1984). The wing beat kinematics anu musculai activity can be
negatively affecteu by the lack of oxygen in these highei elevations. Although
hummingbiius have efficient methous of tianspoiting oxygen in the bouy, this can't
always completely make up foi the lack of physical oxygen in the aii. Bummingbiiu
flight was obseiveu in uiffeient levels of aii uensities (iefeiences 1-S). It was
concluueu that hummingbiius ieach stioke amplituue of 18u uegiees in low-uensity
aii. They ieach a constiaint on flight peifoimance when the stioke amplituue hits
this level. The biius only gieatly inciease theii wing beat fiequency in hypouense
aii. In incieaseu aie uensity anu hypoxia, the hummingbiius mimickeu being in an
enviionment of incieaseu altituue.
NE S2Su Fall 2u1S Teim Papei: Shelby Fostei

In the uecieaseu aii uensity tiials, wing beat fiequency incieaseu as aii uensity
uecieaseu (Figuie A&B). The smallei hummingbiius with smallei wingspans hau a
fastei wing beat than the laigei hummingbiius with the laigei wingspans. The
stioke amplituue incieaseu with aii uensity as well. In the hypoxia tiials, as the
paitial piessuie of oxygen uecieaseu, they showeu a ueciease in wing beat
fiequency, but the oveiall uiffeiences weie slight. The biius stoppeu hoveiing when
the oxygen ieacheu a minimal concentiation, anu then slowly uescenueu to the
flooi. The wing beat fiequency coiielateu moie with the aii uensity iathei than the
level of oxygen. The stioke amplituue showeu no ielationship with the level of
oxygen in the chambei. In the hypeioxia tiials wheie the biius weie given extia
oxygen, no effect was obseiveu on wing beat fiequency oi stioke amplituue.


Concluuing the exploiation of the humming biiu flight mechanism, the
hummingbiiu is oveiall efficient in the way it ueliveis oxygen to the bouy. It has to
be efficient, since it has highei oxygen consuming muscles compaieu to most othei
common veitebiates. They have veiy high mass-specific metabolizing iates. Which
They have small bouies, high enzyme flux capacities, a high iatio of capillaiy suiface
aiea to muscle fibei suiface aiea, anu a laige amount of mitochonuiia to supply
eneigy to the uemanuing muscles in the wings. The size of theii pectoials uemanus
this eneigy, anu uses it foi fast flapping speeus.

NE S2Su Fall 2u1S Teim Papei: Shelby Fostei

The kinematic mechanisms useu foi lift piouuction vaiy with oxygen availability
anu aii uensity. The biius inciease theii lift in hypouense aii mainly by
manipulation of win stioke amplituue, without much change in wing beat fiequency.
A long uuiation of incieaseu wing beat fiequency woulu most likely iequiie much
moie oxygen, but stioke amplituue coulu be sustaineu ovei vaiying levels of aii
uensity conuitions. The hummingbiius aie shown to be moie iesistant to lowei level
of oxygen anu change in elevation. Buiing anaeiobic buists, the biius incieaseu
theii wing beat fiequency by about 2u% (on aveiage). In conclusion, the wing beat
fiequency helps inciease the amount of lift but is limiting baseu on the constiaint of
oxygen ueliveiy to the muscles.

The hummingbiius have poweiful tools foi hoveiing since they can manipulate
many uiffeient mechanisms to aujust to uiffeient enviionmental conuitions. They
can altei theii bouy plane, wing beat fiequency, angulai iotation of the wing, angle
of attack, toision along the wing. They must have evolveu these seveial vaiiables to
suivive low aii uensities anu high levels of elevation. They can withstanu winu gusts
with countei toique causeu by the high speeu flapping of the wings, making it easy
foi them to gathei the nectai fuel that they neeu to sustain the aeiobic activity
necessaiy foi theii unique hoveiing flight. It is not suipiising that they suiviveu the
evolutionaiy piocess, since they aie so efficient in ietiieving the fuel that they neeu
to suivive.


1-Chai, P. Chen, }. S. C. anu Buuley, R. ( 1997). Tiansient hoveiing peifoimance of
hummingbiius unuei conuitions of maximal loauing. }. Exp. Biol. 2uu, 921 -929.

2-Chai, P. anu Buuley, R. ( 199S). Limits to veitebiate locomotei eneigetics
suggesteu by hummingbiius hoveiing in heliox. Natuie S77, 722 -72S.

S-Chai, P. anu Buuley, R. ( 1996). Limits to flight eneigetics of hummingbiius
hoveiing in hypouense anu hypoxic gas mixtuies. }. Exp. Biol. 199, 228S -229S.

4-Chai, P. Baiiykissoon, R. anu Buuley, R. ( 1996). Bummingbiiu hoveiing
peifoimance in hypeioxic heliox: effects of bouy mass anu sex. }. Exp. Biol. 199, 274S

S-Chai, P. anu Nillaiu, B. ( 1997). Flight anu size constiaints: hoveiing peifoimance
of laige hummingbiius unuei maximal loauing. }. Exp. Biol. 2uu, 27S7 -276S.

6-Bickinson, N.B., Lehman, F. 0. anu Sane, S.P. (1999) Wing iotation anu the
aeiouynamic basis
of insect flight, Science, 284(S618): 19S4-196u.

NE S2Su Fall 2u1S Teim Papei: Shelby Fostei
7-Ellington, C. P. ( 1984). The aeiouynamics of hoveiing insect flight. vI. Lift anu
powei iequiiements. Phil. Tians. R. Soc. Lonu. B SuS, 14S -181.

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1u-Lasiewski, R. C. (1964). Bouy tempeiatuie, heait anu bieathing iate anu
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Waiiick B. Chiistophei }. Claik, Bonalu R. Poweis, Tyson L. Beuiick,
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