Lecture 5

Physiology

High body temperature

No matter the environment birds tend to keep their
body temperature around 40 degree Celsius

High body temperature allows for quick reflexes and

movements

Maintenance of high body temperature is extremely costly

Birds consume 20-30 times as much energy as reptiles

Too high (>46 degrees) results in death

In contrast to other animals, birds need quick reflexes, fast
movements, and endurance

Increased aerobic metabolisms make this possible

Increased metabolism demands high rates of energy

and oxygen delivery, as well as rapid removal of
waste products
 Respiratory System
Four anatomic features of the avian respiratory system
nostrils
tracheal system
lungs (relatively small)
air sacs

As opposed to mammals with each breath birds replace nearly all the air in
their lungs.

Respiratory System plays a large role in thermoregulation

Breath through mouth or nostrils (nares)
A flap (Operculum) covers and protects the nostrils (diving and
nectar-feeding birds)

Air enters nasal septum where air is cleaned and warmed

Air enters trachea

Trachae splits at the syrinx

Air enters the lungs

Air sacs provide unidirectional flow of air

Help remove hear
Protect internal organs
So, how does air flow through the avian lungs & air sacs during respiration?

1 - On first inhalation, air flows through the trachea & bronchi & primarily into the
posterior (rear) air sacs
2 - On exhalation, air moves from the posterior air sacs & into the lungs
3 - With the second inhalation, air moves from the lungs & into the anterior
(front) air sacs
4 - With the second exhalation, air moves from the anterior air sacs back into
the trachea & out
 Air Sacs
9 air sacs

Allow unidirectional flow

Have very thin walls

Few blood vessels

Act as bellows to ventilate lungs

Avian lungs

Diagram of parabronchial
Avian tracheas
 Circulatory System
High metabolism of birds demands rapid circulation of high volumes of blood

four-chambered heart

Avian hearts are 50 – 100% for powerful than those of mammals of a

similar size

Legs get 3 times as much blood as the pectoral muscle

The right ventricle pumps blood to the lungs, while the left ventricle pumps blood to the rest of the
body. Because the left ventricle must generate greater pressure to pump blood throughout the body (in
contrast to the right ventricle that pumps blood to the lungs), the walls of the left ventricle are much
thicker & more muscular.

Cross-section of the
ventricles
 Circulatory System
Avian hearts also tend to pump more blood per unit time than mammalian hearts. In other words,
cardiac output (amount of blood pumped per minute) for birds is typically greater than that for mammals
of the same body mass. Cardiac output is influenced by both heart rate (beats per minute) and stroke
volume (blood pumped with each beat). 'Active' birds increase cardiac output primarily by increasing
heart rate. In a pigeon, for example (Butler et al. 1977):

Rest Active Increase

115 670
Heart rate 5.8x
beats/min beats/min
Stroke
1.7 ml 1.59 ml 0.9x
volume
Cardiac
195.5 ml/min 1065 ml/min 5.4x
output
Oxygen
20.3 ml/min 200 ml/min 10x
consumed

In general, bird hearts 'beat' at somewhat lower rates than mammals of the same size but pump more
blood per 'beat.' Among birds, heart rate varies with size
Metabolism
 Metabolism
• Birds have high basal metabolic rates and use energy at high rates. Among birds, songbirds
(passerines) tend to have higher basal metabolic rates than nonpasserines. Hummingbirds, have
the highest basal metabolic rates of all birds. In general, basal metabolic rate (or BMR) is related
to mass, with larger birds expending less energy per unit weight than smaller birds.

• Species Mass (gms) Kcal/kg/day

• Trumpeter Swan 8900 47
• Brown Pelican 3500 75
• Common Raven 850 108
• American Kestrel 110 157
• White-crowned Sparrow 27 324
• House Wren 11 589
• Rufous Hummingbird 3.5 1600
 Heat loss
Birds have no sweat glands so evaporation, one way to lose heat, can occur only via respiratory
system:

gular flutter
rapid fluttering of the gular area
observed in many birds, including pelicans, cormorants, turkey vultures, roadrunners,
quail, & goatsuckers (nighthawks & poor-wills)

Panting

Behavior
Stay in shade
Wet portions of the body

Plumage -
number of feathers varies seasonally, with more during the winter than the summer
position of feathers controlled by dermal muscles
lower temperatures > muscles contract ('goose bumps') to 'erect' feathers >
erect feathers create more air space > more, warm air trapped in the
plumage acts as an effective layer of insulation
 Heat loss

Infrared Thermography image of a House Sparrow exposed to 15°C

and no wind (left) vs. that of a sparrow exposed to 15°C and a wind
speed of 2 meters/second (right). Red colors represent higher
temperatures grading into cooler temperatures with blue colors.
Heat retention
Legs & Feet
heat loss is limited in cold weather because
of a counter-current mechanism that 'saves'
heat can serve as heat 'radiators' during hot
weather: increased blood flow

storks & vultures defecate on legs to increase

heat loss by evaporation
In a countercurrent exchanger, flow in two adjacent tubes (like blood vessels) is in opposite directions. Imagine these are
blood vessels in a bird's leg: the artery on top & the vein on the bottom. The artery is bringing warm blood into the legs. As
the arrows indicate, heat from the blood in the artery is transferred to the blood in the vein (but, of course, oxygen &
nutrients continue on to supply the cells in the feet). As a result of this heat 'exchange', blood in the bird's feet is relatively
cool & little heat is lost. So, even a duck standing on ice loses little heat from its feet.

Birds living in cold environments must conserve body heat to avoid hypothermia. However, blood flowing from the body
core to the periphery (like the legs & feet) carries heat can be readily lost through the skin. To prevent such loss, brids have
a countercurrent heat exchanger - blood vessels in the legs (arteries going in & veins coming out) in close proximity that
allow heat to be recaptured and saved. The principle of countercurrent heat exchange is so effective and ingenious that it
has also been adapted in human engineering projects to avoid energy waste, e.g., by ensuring good ventilation of buildings
while avoiding the loss of heat to the environment on a cold winter's day.
 Cold stress

At and below 0 degrees C, intermittent pulses of increased blood flow to

extremities are used to prevent freezing and tissue damage, reflected in
increased heat loss from the feet and an increased metabolic rate
(Kilgore and Schmidt-Nielsen 1975).
 Cold Stress
• hypothermia is generally triggered by reduced
availability of food or depletion of stored energy
reserves or both
• two major categories of hypothermia in birds:
• nocturnal hypothermia:
– body temperature may decline 8 - 10 degrees (no
lower than about 30 degrees C)
– observed in doves & pigeons, turkey vultures, &
several passerines, including chickadees
– conserves energy to help a bird survive periods of
reduced food availability or, in some cases, to
facilitate the process of premigratory 'fattening'
 Stop heat loss
Posture at low temperatures
birds can withdraw feet into plumage to reduce heat
loss

tuck head & neck under wing to reduce heat loss

 Stop heat loss
Communal roosts and huddling can also
conserve heat

100 nuthatches in a cavity

3 kinglets on a branch

Inca doves forming a dove pyramid

Ptarmigans and snow bunting borrow into the snow

 Huddling behaviour.
• E.g. Emperor penguin.
• Male incubates egg on feet ~3 months.
• Huddling penguins lose only half fat per day as
when alone.
• Temp. in huddle can reach 35C.
• Animals exposed on windward
side continually move leeward.
• Huddling also seen with chicks
in bird nests. More energy for
growth not heat production.
During the winter, Willow Ptarmigans produce a denser coat of feathers and actually lower
basal metabolic rates, which reduces the gradient between the internal body temperature and
the external temperature. This is the same as keeping your house at a cooler temperature to
reduce your heating bill. Both responses reduce the amount of energy necessary to stay warm.
 Torpor in the hummingbird.

50
O2 consumption
(ml O2/g/h)

Dawn Noon Dusk

 Torpor
• Big Birds Can Keep Their Cool Too - Researchers have found the
biggest example yet of a bird that cools down to conserve energy
(Kortner et al. 2001). The Australian Tawny Frogmouth (Podargus
strigoides), which weighs in at half a kilogram, uses ‘torpor’ — as a
controlled reduction of body temperature is properly called — as a
survival strategy in winter, even though it is almost ten times larger
than any other bird known to do so. Fritz Geiser and colleagues of
the University of New England, Armidale, Australia, fitted seven
frogmouths with external temperature-sensitive transmitters and
three of those with internal temperature-sensitive transmitters, and
tracked the birds’ temperatures every 10 minutes for 9 months.
During winter, Geiser’s team was surprised to find, these big birds
regularly enter torpor. “This enables the bird to remain resident in its
territory throughout the year.” Avian torpor, they predict, is much
more common than is currently believed.
Acclimation

Seasonal variation in summit metabolism and

cold tolerance for American Goldfinches. Bars
represent the mean ± SD summit metabolism achieved
by acclimatized American Goldfinches in winter
(Jan–Feb), spring, and summer (June–August).
Mean ± SD temperature at cold-limit (Tcl), the
bath temperature at which goldfinches became
hypothermic, is represented by circles connected
by a line. Note the positive relationship between
summit metabolism and cold tolerance in this
species
 Acclimation
Seasonal acclimatization by American Goldfinches -- Liknes et al. (2002)
evaluated seasonal changes in cold tolerance, basal metabolic rate (BMR), and
summit metabolic rate (maximum rate of metabolism in response to cold exposure)
for American Goldfinches (Carduelis tristis) from South Dakota to determine if
goldfinches differ in pattern of metabolic acclimatization from other species of small
birds. Goldfinches were captured in winter (Jan–Feb), spring (April), and summer
(June–August) and tested on the day of capture. Cold exposure tests involved
subjecting individual birds to a decreasing series of temperature. The temperature
eliciting hypothermia was designated the cold limit (Tcl). Whole-animal metabolic
rates were analyzed. Winter goldfinches demonstrated significantly higher BMR
(46%) and summit metabolic rates (31%) and significantly lower Tcl (- 9.5°C vs.
1.3°C) than their summer counterparts. Spring goldfinches also showed significantly
higher summit metabolic rates (21%) and significantly lower Tcl (- 5.3°C) than
summer birds. Winter birds had higher BMR (23%) and summit metabolic rates (8%)
than spring birds. In winter birds, Tcl was also significantly lower than in spring birds.
These data support the view that prominent winter increases in summit metabolic
rates and BMR are components of winter acclimatization in American Goldfinches in
South Dakota and that seasonal changes in metabolism in goldfinches are similar to
those for other small temperate-wintering birds
 Bergmann’s Rule
• The increase in body size with cooler
temperatures

Bergmann's Rule in Downy Woodpeckers (Picoides pubescens)

"A north-south cline in wing length, a measure of body size .... Each line
connects populations with the same mean wing length (in mm)." (from
Futuyma 1998 after James 1970)
 Water stress
• Because water is a by-product of metabolism and birds have high
metabolisms they tend to not have to drink too much water

• Some birds drink while flying (swallows)

• In arid environments thousands of birds will decent on water holes

• In mesic environments drinking at a water hole is rare

 Excretory system
• The most obvious adaptation to conserving water is the excretion of
nitrogenous waster in the form of uric acid. These white crystals
give bird droppings their usual white color.

• Birds only need 1/20th of the water to excrete the same amount of
uric acid as mammals

• Bird excrement can be anywhere from 90% water to 55% water in

arid environments
 Excretory system
• A problem for oceanic birds is the salt in the water they ingest

• Seabirds as well as a few other species have salt glands

• Salt glands excrete salt rapidly after being ingested

• A gull can excrete 90% of the salt in the water it drinks within 3
hours

• Some birds have more developed salt glands than others

 Energy Budget of migrants
• Wikelski et al. (2003) found that migrating
thrushes use more energy during stopovers
than on the wing.

• . Finding food during rest breaks and sitting

through the cold night can use lots of
energy.

• Every spring, millions of Catharus thrushes

migrate from Panama to Canada - a journey
of 4,800 km that takes around 40 days.

• They have regular breaks and fly for just a

few hours at night.

• Over the entire journey, the energy used by

a single bird is equivalent to that in half a
kilo of worms. Less then 30% of this is used
on flight, the team found.

• As the birds headed north, the team gave

chase in a fleet of cars and a light airplane.
"This was the easy part," says Visser. The
hard bit was catching them at the other end,
he recalls. - Helen R. Pilcher, Nature
Science Update
 Dominance and energy
• Dominant birds stay leaner than their subordinates

• Legend says that the early bird gets the worm, but research
suggests that the bird that dines just before going to bed has
the real advantage.

• Pravosudov et al. (1999) found that socially dominant birds

in three species (Carolina Chickadee, Tufted Titmouse, and
White-breasted Nuthatch) are generally leaner than
subordinate peers, probably because they can eat
when they want.

• Dominant birds stay lean during the day, then pack on the fat just before a
chilly winter night.

• Staying lean helps birds stay more maneuverable during attacks by predators.
Lean birds also have more time to watch for predators, rather than looking for
food.

• Subordinate birds must carry more fat during the day as an 'insurance policy
making them more vulnerable to capture by predators

• A bird can gain as much as 10% of its total body mass each day in fat. Gaining
fat before nightfall can help birds survive in winter because they often go into
hypothermia as a survival mechanism.

• Extra fat at roosting time means a bird needs to go less far into hypothermia at
night because it has more energy for metabolism. Hypothermia is thought to
be a cost, because it makes a bird less aware of its surroundings, therefore
increasing its vulnerability to nocturnal predators.