Feathers and Flight

Feathers
Primary function
- insulation
- aerodynamics
- communication
- camouflage

Secondary function
- swimming
- sound production
- hearing
- protection
- cleanliness
- water proofing
- water transport
- tactile sensation
- support
Lecture 4

Feathers and Flight

Keratin – an inert substance of insoluble microscopic filaments embedded
in an amorphous protein matrix

Types of feathers
Contour – body feathers

Down - insulation

Semiplume – insulation, aerodynamics, courtship

Filoplume – aerodynamic adjustments (sensory)

Bristles – sensory and protection

Powder down – waterproofing

• While most other birds' feathers produce a
small amount of powder, powder feathers
(yellow feathers near the bottom of the photo
that form a "V") are highly developed in herons,
egrets, & bitterns and the quill of the powder
down is continuously growing and
disintegrating, thus creating the powdery
substance. The barbs of powder feathers
disintegrate, providing a fine powder that is
thought to aid in preening and waterproofing
the other feathers. They are the only feathers
that grow continuously and are never molted.
While many species have powder feathers
scattered within patches of normal down, they
are most prominent in the herons and bitterns
(family Ardeidae), and are located on the
breast and belly.
 Capital tract

Ventral tract

Spinal tract

Alar tract

Humeral tract

Ventral tract

Femoral tract

Crucal tract Caudal tract

Feathers and Flight
 Feature Care

Tundra Swans have ~25,000 feathers of which 20,000 are on the head

Uropygial gland (preen gland) – this gland on the rump of most birds
secrets a rich oil of waxes, fatty acids, fat, and water.

Preening – birds use there bill to apply these substances to feathers in

order to clean and preserve them. The substances also help insulate and
waterproof the birds.

Feather lice – reduce the structural integrity of feathers.

Hippobascid flies

Bills and feat are important in removing lice

Some birds are poisonous (New Guinea shrike-thrushes)

 Feature Growth

Feathers grow from specialized pockets of cells called follicles

Feathers are held in place by basal cells called calamus

Feathers can be tough to pull out

Feathers can fall out due to fear (fright molt)

 Color pigments in feathers

• Melanins: Dark, brick-red, tan, brown

• Carotenoids: From diet, bright red, yellow, orange

• Porphyrins: from hemoglobin, green, red, brown

• Structural: prismatic; scattered, reflective, melanin in

barbules.
• Avian Plumage Color (Prum et al. 2003) -- The colors of avian plumage are produced
by chemical pigments (e.g., melanin or carotenoids) or by nanometer-scale biological
structures that differentially scatter, or reflect, wavelengths of light. No exclusively blue
or UV-colored pigments are known in vertebrates, but various carotenoid pigments in
bird feathers produce UV wavelengths in combination with human-visible yellow,
orange, or red colors. Ultraviolet structural colors of feathers can be produced by two
types of structures. Primarily iridescent colors are produced by arrays of melanin
granules in feather barbules. Those structural colors are created by coherent
scattering, or constructive interference, of light waves scattered from the layers of
melanin granules in barbules. A few species of hummingbirds and European Starlings
are known to produce UV hues with coherently scattering melanin arrays in feather
barbules. The most commonly distributed UV hues, however, are structural colors
produced by light scattering from the spongy medullary layer of feather barbs. To date,
primarily UV hues have been documented in the feather barbs of Chalcopsitta
cockatoos (Psittacidae) and Myiophonus thrushes (Turdidae). Extensively UV hues
with a peak reflectance in the human-visible blue range have been observed in feather
barbs of Blue Tits (Parus caeruleus), Bluethroats (Luscinia svecica), and Blue
Grosbeak. In addition, Prum et al. (2003) have found extensive UV reflectance from
apparently blue feather barbs in many families and orders of birds including motmots
(Momotidae), manakins (Pipridae), cotingas (Cotingidae), fairy wrens (Maluridae),
bluebirds (Sialia), buntings and others. The structural UV hues of feather barbs, like
other barb structural colors, are produced by the keratin air matrix of the spongy
medullary layer of the barb ramus. However, the precise physical mechanism by which
the human-visible and UV barb colors are produced remains controversial. Analysis of
the spongy medullary keratin of UV-colored feather barbs of Myiophonus caerulea by
Prum et al. (2003) demonstrated that, in this species, color-producing tissue is
substantially nanostructured at the appropriate spatial scale to produce the observed
ultraviolet hues by coherent scattering, or constructive interference.
Transmission electron micrographs of the spongy medullary keratin of the UV-colored feather barbs of
Myiophonus caeruleus.

Left panel: cross-section of a UV-colored feather barb ramus showing the solid keratin of the barb
cortex at the periphery, three adjacent medullary cells with spongy keratin matrix and cell walls,
and melanosomes around the large vacuole at the center of the barb ramus. Right panel: close-up of
the spongy medullary matrix of keratin bars and air vacuoles. Scale bar equals 500 nm. Abbreviations:
c = barb cortex; cw = cell wall, k = spongy medullary keratin, m = melanosome, and v = air-filled
vacuole
The blue grosbeak is one of the best examples of a bird that utilizes the
ultraviolet spectrum. While female grosbeaks are brown, male grosbeaks are a
brilliant blue. The brilliant blue comes from the males' feather structure; after
measuring peak feather wavelength (when the most light is reflected) and
intensity under a spectrophometer, both peak wavelength and intensity were
found to fall in the ultraviolet range. The intensity of UV reflectance varies from
male to male, so female grosbeaks can tell a difference while we humans
cannot
 Melanin

Why do white birds have black wing tips?

What is great about Melanin
Melanin strengthens feather

Gloger’s rule – wet climate tend to have dark colored birds

Melanin may promote drying of damp feathers by absorbing and

concentrating radiant hear in the feather microstructure

If there are more ectoparasites in warm environments melanin

may help protect the feathers from these parasites
House Finches

Geoff Hill’s Lab Auburn University

• Whistle through the wing - Birds molt for a variety of reasons. Molting regulates body
temperature, keeps feathers neat and waterproof and allows seasonal changes in
appearance for mating or migration. However, generating new feathers uses extra
energy; staying warm with less plumage uses extra energy, and flying with smaller,
work-in-progress wings requires extra energy. So, not all birds molt in the same way.
Ducks, swans and geese, for example, shed all their flight feathers at once and are
flightless until replacements have grown. Most other birds, however, lose and renew
their feathers according to a continuous, pre-programmed sequence. This sequential
molting gives rise to a range of temporary feather gaps that seem to reduce take-off
speed, take-off angle and level flight speed and to impede predator evasion by raising
a bird's minimum turning radius. Anders Hedenstrom and Shigeru Sunada of
Cambridge University estimated how the aerodynamics of flight are affected by molting
(Hedenstrom & Sunada 1999). They estimated drag and lift by analyzing the fluid
dynamics of symmetrical gaps in flat, rectangular model wings of various width-to-
length (aspect) ratios, at a fixed angle with respect to air flow – a system that
reasonably approximates a bird in gliding, but not flapping, flight. Although the effects
were small, Hedenstrom and Sunada concluded that both feather gap size and position
affect flight performance. Large gaps, and gaps in the middle of the wing impede
aerodynamic efficiency more than small, wing-tip gaps. They also found that the
detrimental effect of molt gaps increases with increasing aspect ratio. In other words a
bird with short, broad wings, like a vulture, won't miss a few feathers as much as one
with long, narrow wings, like an albatross. "This is of great ecological significance," they
muse, "as it could help explain why large birds show relatively slow rates of molting that
are associated with rather small gaps." -- Sara Abdulla, Nature Science Update
 Color phases

Opposites attract. Two color morphs

occur with equal frequency among
White-throated Sparrows, and each
morph nearly always mates with the
other. "White-stripe" morphs have
sharply contrasting black-and-white
head stripes and usually solid gray
breasts; "tan-stripe" morphs have less-
contrasting dull black-and-tan colored
stripes, and they often have streaks on
their breasts. The more aggressive
"white-stripe" females are preferred by
both forms and are more successful in
attracting the preferred "tan-stripe"
males. This combination forms most
rapidly, leaving "white-stripe" males
and "tan-stripe" females to pair with
each other. The result is an observed
96 percent frequency of mixed-morph
pairings
 Flight

Schematic cross-section through a bird bone.

A - periosteal surface
B - lamellar cortical layer,
C - endiosteal surface
D - trabecular layer,
E - pores/pneumatic openings/blood vessel openings
(From: Davis 1998)

Bird bones are pneumatic (filled with air spaces but reinforced with internal
struts or trabeculae)
large, soaring birds (e.g., albatrosses, vultures, & hawks) have
pneumatic bones
divers (e.g., loons) must be 'heavier' to dive & so do not have
pneumatic bones
small birds (e.g., many passerines) do not have pneumatic bones (their bones
are so small that little weight would be 'saved' with pneumatic bones)
Modification for flight
thoracic region:
tightly articulated vertebrae
ribs with upper (vertebral) & lower (sternal) segments that connect the vertebral column
with the sternum
uncinate processes overlap successive ribs to reinforce rib-cage

pelvic region:
pelvic (innominate) bones are fused with lumbar & sacral vertebrae. The fused portion of
the vertebral column is called the synsacrum & it's composed of the last thoracic
vertebra, the lumbars, sacrals, & anterior caudals.
last few caudal vertebrae are partially fused to form the pygostyle (that helps support the
tail feathers)
ulna (the bone that supports the secondaries) is enlarged

carpal bones (wrist bones) are reduced in number (just 2)

metacarpals (palm bones) - 1st & 5th metacarpals are

lost; 2nd, 3rd, & 4th are united (with vestigial carpals) to
form the carpometacarpus

digits - only 3 (rather than the typical 5 found in most

vertebrates)

phalanges (the bones that make up the digits) - few in

number; 4 - 7 make up the 3 digits
Keel or sternum is a large bone for the attachment of the large mussels
associated with flight

No keel in flightless birds

Tendon of the supracoracoideus passing through the
foramen triosseum and inserting on the humerus
(From: Degernes and Feduccia 2001)
jaw muscles are reduced in many birds (powerful muscles often unnecessary
because food is swallowed whole or in large pieces, e.g., owls)

hindlimb muscles reduced in many species because:

the rigid skeleton of birds (hindlimb, pelvic girdle, & synsacrum) provides
much support &, as a result, less musculature is needed

hindlimbs are sometimes used for little else but perching (e.g., hummingbirds
& swifts
Why does the slower moving air generate more pressure against the wing than the faster moving air? In calm air,
the molecules are moving randomly in all directions. However, when air begins to move, most (but not all) molecules are
moving in the same direction. The faster the air moves, the greater the number of air molecules moving in the same
direction. So, air moving a bit slower will have more molecules moving in other directions. In the case of a wing,
because air under the wing is moving a bit slower than air over the wing, more air molecules will be striking the bottom
of the wing than will be striking the top of the wing. This is called the Bernoulli effect & this creates lift!
Wings also provide lift through Newton's Third Law of Motion which states that for every action there is an equal and
opposite reaction. As the wing moves though the air, the lower surface of the wing deflects some of the air downward.
As Newton's Third Law of Motion explains, an additional force is generated. The deflected airflow underneath the wing
is the action. The reaction is that the wing moves in the opposite direction (in this case, upwards). This means that the
development of low pressure above the wing (Bernoulli's Principle) and the wing's reaction to the deflected air
underneath it (Newton's third Law) both contribute to the total lift force generated.

When the curvature over the top becomes greater by increasing the angle of attack (below), the air moves even
faster over the top of the wing and more lift is generated. Eventually, however, if the angle of attack becomes too great,
the flow separates off the wing and less lift is generated. The result is stalling. For birds, the optimum angle of attack
is typically about 3 - 5 degrees. Birds also tend to stall at low speeds because slower moving air may not move
smoothly over the wing
Soaring & thermals
How do gulls hang in the air?
 Alula

At low speeds (such as during take-off & landing), birds can maintain smooth
air flow over the wing (and, therefore, maintain lift) by using the alula. The
alula is formed by feathers (usually 3 or 4) attached to the first digit
 Landings

Birds are the only animals that are able to land perfectly on their hindlimbs
Hovering is hard work for most birds - Ever seen a songbird hover over a crowded feeding station,
waiting for a perch to open up so it can land and eat? Looks like hard work, doesn't it? It is, which is
why hovering is something most birds don't like to do -- or can't do -- for very long. Kenneth P. Dial of
the University of Montana and colleagues (Dial et al. 1997) surgically implanted strain gauges in the
wings of three Black-billed Magpies. The devices measured the force exerted by the main flapping
muscle with each wing beat. The birds then flew in a wind tunnel at a range of speeds. The strain
gauge allowed the scientists to calculate the power (the amount of work done per unit time) required
to maintain a given speed. Hovering took nearly twice as much power as flying at average speed,
the researchers found. Even when the magpies flew at top speed, they expended far less power than
they did when they hovered. Evidence suggested that when they hovered, the birds were working at
their physical limits. Their wing muscles appeared to be employing anaerobic metabolism, a source of
energy that can't be sustained for long. There are clearly exceptions to this. Hummingbirds, the
authors note, have an unusual shoulder design that allows them to generate lift on both down-beat
and up-beat. But birds with a body design similar to magpies are likely to have strict limits on their
abilities to fly standing still.
 Why fly in a “v”
Birds conserve fuel by flying in V formations. By measuring heart rates, researchers in France now
have proof that pelicans use 11-14% less energy flying together, even when they are not perfectly
positioned to take advantage of the wake from those in front of them. Configured flight may create a
stream of air that allows birds to glide longer, suggests Henri Weimerskirch, the biologist at the National
Centre of Scientific Research at Villiers en Bois, who led the study. "If you look closely, you see that the
birds at the back are gliding more than the leader." People have been asking whether V formations are
more efficient for more than 100 years, Speakman says, but no one had measured energy savings
before. "They took a century-old problem and went to the heart of it," he says.

Gliding not flapping

Scientists from the Centre National de la Recherche Scientifique, Villiers en Bois, France, taped heart
monitors to the birds' backs. The pelicans had been trained to follow a light aircraft and a motor boat,
allowing the team to observe and study them during flight. The researchers found that when the
pelicans were flying in formation - benefiting from each other's airstreams - their heart rates were lower
than when they were flying solo. They also spent more time gliding. "They fly in formation to save
energy," team leader Henri Weimerskirch told BBC News Online. "It's not because they are using the
upward airstream of their neighbour, it is because they are able to glide more often."

Aerodynamic models
The aerodynamic benefit of formation flying has long been suspected. But until now, it has proved
impossible to test. "Everything that has been done until now has been theoretical, based on
aerodynamic models," said Dr Weimerskirch. "It cannot take in the full complexity of the flight of the
bird." The work, published in the journal Nature, suggests that formation flight evolved because it
allowed birds to reduce their energy expenditure and fly further. This would be an advantage both for
migration and in hunting for food. Flying in a "V" also has social advantages: it allows birds to
communicate with each other while on the wing.
Why fly in a “v”
Flightless birds