1) Hummingbirds have evolved unique flight abilities that allow them to hover and extract nectar from flowers efficiently as their main food source. Their anatomy supports high-frequency wingbeats through adaptations like large pectoral muscles and high mitochondrial content in flight muscles.
2) Their small size and metabolism allow hummingbirds to achieve wingbeat frequencies as high as 80 beats per second during hovering, expending much more energy than other birds. Flower nectar is very energy-dense, fueling their demanding flight.
3) Studies show hummingbird aerodynamics are similar to insects, producing lift on both upstrokes and downstrokes through wing rotation. However, their anatomy and kinematics are still
1) Hummingbirds have evolved unique flight abilities that allow them to hover and extract nectar from flowers efficiently as their main food source. Their anatomy supports high-frequency wingbeats through adaptations like large pectoral muscles and high mitochondrial content in flight muscles.
2) Their small size and metabolism allow hummingbirds to achieve wingbeat frequencies as high as 80 beats per second during hovering, expending much more energy than other birds. Flower nectar is very energy-dense, fueling their demanding flight.
3) Studies show hummingbird aerodynamics are similar to insects, producing lift on both upstrokes and downstrokes through wing rotation. However, their anatomy and kinematics are still
1) Hummingbirds have evolved unique flight abilities that allow them to hover and extract nectar from flowers efficiently as their main food source. Their anatomy supports high-frequency wingbeats through adaptations like large pectoral muscles and high mitochondrial content in flight muscles.
2) Their small size and metabolism allow hummingbirds to achieve wingbeat frequencies as high as 80 beats per second during hovering, expending much more energy than other birds. Flower nectar is very energy-dense, fueling their demanding flight.
3) Studies show hummingbird aerodynamics are similar to insects, producing lift on both upstrokes and downstrokes through wing rotation. However, their anatomy and kinematics are still
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 iecoiueu.
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.
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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.
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