ANSC 3180 Lecture Notes - Lecture 7: Terrestrial Locomotion, Chukar Partridge, Passerine
2 May 2018
School
Department
Course
Professor
Metabolizable energy (ME) -intake at which body weight and composition
remain constant, in a healthy, non-reproducing animal/bird living in its
normal environment
○
Daily energy expenditure (DEE) = BMR + activity + thermoregulation +
food metabolism
○
Maintenance energy requirements:
•
Terrestrial mammals (eutherians) -2.3 x BMR
○
Small, insectivorous marsupials -6.3 x BMR
○
Large, herbivorous marsupials -2.5 x BMR
○
Passerine birds -2.8 x BMR
○
Non-passerine birds -2.5 x BMR
○
Seabirds at sea -3.3 x BMR
○
Birds feeding young ->4.0 x BMR
○
Hummingbird feeding young -8.0 x BMR
○
Nesting seabirds -2.1 x BMR
○
Daily energy expenditure (free-ranging):
•
It is not the 'average' but the 'variation' that determines whether
animals reproduce, survive or die
!
*these values are only generalizations/ averages
Maintenance Energy: Summary
Measure
Basal
Metabolism
Nutrient
Metabolism*
Activity
Thermoregula
tion
Basal (BMR)
X
Standard (SMR)
X
(X)
Fasting (FMR)
X
X
Resting (RMR)
X
X
X
Average Daily
(ADMR)
X
X
X
X
Existence
Metabolism (EM)
X
X
X
X
Existence Energy
(EE)
X
X
X
X
Metabolized Energy
(ME)
X X X X
Field (FMR) X X X X
Maintenance Energy/ Daily Energy Expenditure -Measures of Metabolism
*Nutrient/Food Metabolism = Heat Increment of Feeding (HI) = Specific Dynamic
Action (SDA)
Exceptions: equids (horse family), passively lock legs to remain
standing (elephants)
!
Requires energy, ~20% greater than lying (in mammals)
○
Painted quail, bobwhite quail, chukar, partride, guinea fowl, wild
turkey, domestic chicken
!
Birds, estimated at 17 +/-11 % greater than sitting
○
Standing
•
Walking, trotting, running
○
Energy used increases linearly with speed
○
Energy cost of increasing speed of 1km/ hr is greater in smaller
animals
!
Energy cost (kcal/kg/hr) for a given speed is higher for smaller animals
○
*cheetah: ~110 km/h
○
Increasing energy use remains basically linear, when energy
cost of walking at a faster speed is higher than trotting at the
same higher speed
□
Ex. Elk calk changing from a walk -trot -gallop
!
E.g. hopping in kangaroos, bounding in mink
□
Exceptions: changing to a faster gait decreases energy used, more
efficient/ unit distance covered
!
Gait may change when the energy cost of increasing speed using the slower
gait is greater than changing to a faster gait
○
Y -kcal/kg/km
□
X -body weight (kg)
□
Y = 2.5 X-0.316 *for both quadrupeds and bipeds
!
Exceptions: penguins/ waterfowl waddling (twice as much energy is
required)
Y = 5.57 X-0.327
!
Energy cost of locomotion (kcal/kg/km) is highest in small animals
!
Longer stride length
□
Less frequent strides
□
Body and leg length increases with increasing body weight (larger
animals):
!
Net locomotion cost decreases as young animals grow
□
Net locomotion cost: short-legged domestic reindeer (0.85
kcal/kg/km) is approx. twice the energy cost compared to longer-
legged caribou (0.41 kcal/kg/km) although they have similar body
weights
!
Energy used in horizontal locomotion:
○
0.5% (10g mammal) -4.2% (100kg mammal)
!
Exceptions: migration, locomotion through snow (deer, elk,
caribou, bighorn sheep, penguin)
□
Normally, not a major energy expense
!
Increasing snow depth
!
Increasing snow density
!
Energy costs increase with…
□
Some raise legs higher and then 'bound' (mule deer,
white-tailed deer, caribou, elk, bighorn sheep)
!
'plowing' requires less energy if snow is powdery;
'bounding' involves vertical movement
!
Breaking through snow with surface ice crust involves
vertical movement (extra energy)
!
Following a 'lead' animal through snow reduces energy
costs
!
Some species 'plow' through snow (bison, mountain goats)
□
Light foot-loading: caribou, wolves
◊
Intermediate food-loading: mule deer, white-tailed
deer, moutain goats, bighorn sheep
◊
Heavy foot loading: bison, moose, pronghorns, elk
◊
Young animals have lighter food-loading than
adults
◊
Big feet, small body weight = low energy to move
!
Foot-loading (weight/unit area) and foot area are important in
energy cost of locomotion in snow
□
Caribou run from wolves (longer legs, same loading)
!
Moose stand and fight (longer legs, heavier foot-loading,
equivalent energy costs in snow)
!
Wolves & prey species in snow:
□
'toboggan' in soft snow -waddling is high energy cost
!
'toboggan' on ice -waddling on slippery surface is
difficult
!
But increased feather wear while 'tobogganing' --> may
waddle on hard snow
!
Penguin locomotion in snow:
□
Mammal locomotion in snow:
!
Locomotion as part of daily energy expenditure:
○
Terrestrial Locomotion -horizontal
•
In some cases, descending a slope may require more energy than
moving the same distance horizontally (ex. Bighorn sheep on cliff)
!
Energy required, ~6kcal/kg/vertical km *for all species, body weights and
angles of ascent
○
Terrestrial locomotion -vertical
•
Energy required, 1.6-5.8 x Resting MR (rodents, insectivorous moles)
○
Scraping soil from tunnel walls
!
Removing scraped soil
!
Burrowing includes:
○
Pocket gophers (burrowing in clay soil) requires ~8x energy
compared to burrowing in sandy soil
!
Energy depends on soil type:
○
Access to food
!
Protection from predators
!
A more stable thermal environment
!
Burrowing required 360-3400x energy costs than walking, BUT supplies:
○
Burrowing
•
Energy required is 3.4 x BMR
!
Ex. Herring gull, wandering albatross
!
Gliding flight:
○
Large wings compared to body weight -extremely aerial species
!
Energy required is 3.4-6.4 x BMR
!
Ex. Purple martin, house martin, barn swallow, bank swallow, sooty
tern
!
Very active flight:
○
Without the large wings of extremely aerial species
!
Energy required is up to 14 x BMR
!
Active Flight:
○
Ex. European robin: ~23 x BMR
!
For very short flights (few seconds), energy requirements are even higher
○
Energy costs of flight in bats is very similar to that of birds
○
Ex. 10g bird flying 1km has <1% cost compared to 10g mouse
walking/running 1km
!
Energy costs are high/unit time but low/unit distance
○
Flying
•
Less energy required compared to running or flying (per unit distance) in
terrestrial mammals and birds
○
Most energy efficient speed (per unit distance) is ~0.5 m/s (for most
aquatic mammals and birds)
!
Surface swimming -energy required increases with speed
○
~0.6 m/s for 1kg animal
□
~1.3 m/s for 60kg animal
□
Most efficient speed:
!
Submerged swimming -more energy efficient at faster speeds (less
drag/resistance)
○
~2 x RMR (captive animals)
!
~6-9.3 x RMR in wild penguins catching prey
!
Energy required when swimming at most energy efficient speeds:
○
Energy for high speed swimming to catch prey is up to 50 x
RMR
□
Energy cost is too high --> starvation
□
Sea lion (45kg) with fishing net (0.6kg): energy swimming is 4.3x
higher with net
!
Energy costs of swimming are greatly increased when fishing net, plastic
buoys, or garbage become attached to marine mammal or bird
○
Swimming
•
Swinging from tree to tree using arms
○
Ex. Spider monkey -x1.5 normal walking
!
Energy cost (kcal/kg/km)
○
Spider monkey/slow loris energy use increases ~65% compared to
resting
!
Hanging without movement:
○
3x more energy than 20% required by most standing animals compared to
lying
○
Brachiation
•
Energy required to capture (predators), manipulate, chew and ingest food
○
Ruminants: 20-59% increase over standing
○
Birds: ~11% increase over perching
○
Feeding
•
Fighting (hierarchy and mating rights)
○
Playing
○
Grooming
○
Ruminating
○
Escaping from predators
○
Other Activities:
•
Maintenance Energy -Activity
Energy: Maintenance & Activity
Monday,+February+ 6,+2017
11:37+AM
Metabolizable energy (ME) -intake at which body weight and composition
remain constant, in a healthy, non-reproducing animal/bird living in its
normal environment
○
Daily energy expenditure (DEE) = BMR + activity + thermoregulation +
food metabolism
○
Maintenance energy requirements:
•
Terrestrial mammals (eutherians) -2.3 x BMR
○
Small, insectivorous marsupials -6.3 x BMR
○
Large, herbivorous marsupials -2.5 x BMR
○
Passerine birds -2.8 x BMR
○
Non-passerine birds -2.5 x BMR
○
Seabirds at sea -3.3 x BMR
○
Birds feeding young ->4.0 x BMR
○
Hummingbird feeding young -8.0 x BMR
○
Nesting seabirds -2.1 x BMR
○
Daily energy expenditure (free-ranging):
•
It is not the 'average' but the 'variation' that determines whether
animals reproduce, survive or die
!
*these values are only generalizations/ averages
Maintenance Energy: Summary
Measure Basal
Metabolism
Nutrient
Metabolism*
Activity Thermoregula
tion
Basal (BMR) X
Standard (SMR) X(X)
Fasting (FMR) X X
Resting (RMR) X X X
Average Daily
(ADMR)
X X X X
Existence
Metabolism (EM)
X X X X
Existence Energy
(EE)
X
X
X
X
Metabolized Energy
(ME)
X
X
X
X
Field (FMR)
X
X
X
X
Maintenance Energy/ Daily Energy Expenditure -Measures of Metabolism
*Nutrient/Food Metabolism = Heat Increment of Feeding (HI) = Specific Dynamic
Action (SDA)
Exceptions: equids (horse family), passively lock legs to remain
standing (elephants)
!
Requires energy, ~20% greater than lying (in mammals)
○
Painted quail, bobwhite quail, chukar, partride, guinea fowl, wild
turkey, domestic chicken
!
Birds, estimated at 17 +/-11 % greater than sitting
○
Standing
•
Walking, trotting, running
○
Energy used increases linearly with speed
○
Energy cost of increasing speed of 1km/ hr is greater in smaller
animals
!
Energy cost (kcal/kg/hr) for a given speed is higher for smaller animals
○
*cheetah: ~110 km/h
○
Increasing energy use remains basically linear, when energy
cost of walking at a faster speed is higher than trotting at the
same higher speed
□
Ex. Elk calk changing from a walk -trot -gallop
!
E.g. hopping in kangaroos, bounding in mink
□
Exceptions: changing to a faster gait decreases energy used, more
efficient/ unit distance covered
!
Gait may change when the energy cost of increasing speed using the slower
gait is greater than changing to a faster gait
○
Y -kcal/kg/km
□
X -body weight (kg)
□
Y = 2.5 X-0.316 *for both quadrupeds and bipeds
!
Exceptions: penguins/ waterfowl waddling (twice as much energy is
required)
Y = 5.57 X-0.327
!
Energy cost of locomotion (kcal/kg/km) is highest in small animals
!
Longer stride length
□
Less frequent strides
□
Body and leg length increases with increasing body weight (larger
animals):
!
Net locomotion cost decreases as young animals grow
□
Net locomotion cost: short-legged domestic reindeer (0.85
kcal/kg/km) is approx. twice the energy cost compared to longer-
legged caribou (0.41 kcal/kg/km) although they have similar body
weights
!
Energy used in horizontal locomotion:
○
0.5% (10g mammal) -4.2% (100kg mammal)
!
Exceptions: migration, locomotion through snow (deer, elk,
caribou, bighorn sheep, penguin)
□
Normally, not a major energy expense
!
Increasing snow depth
!
Increasing snow density
!
Energy costs increase with…
□
Some raise legs higher and then 'bound' (mule deer,
white-tailed deer, caribou, elk, bighorn sheep)
!
'plowing' requires less energy if snow is powdery;
'bounding' involves vertical movement
!
Breaking through snow with surface ice crust involves
vertical movement (extra energy)
!
Following a 'lead' animal through snow reduces energy
costs
!
Some species 'plow' through snow (bison, mountain goats)
□
Light foot-loading: caribou, wolves
◊
Intermediate food-loading: mule deer, white-tailed
deer, moutain goats, bighorn sheep
◊
Heavy foot loading: bison, moose, pronghorns, elk
◊
Young animals have lighter food-loading than
adults
◊
Big feet, small body weight = low energy to move
!
Foot-loading (weight/unit area) and foot area are important in
energy cost of locomotion in snow
□
Caribou run from wolves (longer legs, same loading)
!
Moose stand and fight (longer legs, heavier foot-loading,
equivalent energy costs in snow)
!
Wolves & prey species in snow:
□
'toboggan' in soft snow -waddling is high energy cost
!
'toboggan' on ice -waddling on slippery surface is
difficult
!
But increased feather wear while 'tobogganing' --> may
waddle on hard snow
!
Penguin locomotion in snow:
□
Mammal locomotion in snow:
!
Locomotion as part of daily energy expenditure:
○
Terrestrial Locomotion -horizontal
•
In some cases, descending a slope may require more energy than
moving the same distance horizontally (ex. Bighorn sheep on cliff)
!
Energy required, ~6kcal/kg/vertical km *for all species, body weights and
angles of ascent
○
Terrestrial locomotion -vertical
•
Energy required, 1.6-5.8 x Resting MR (rodents, insectivorous moles)
○
Scraping soil from tunnel walls
!
Removing scraped soil
!
Burrowing includes:
○
Pocket gophers (burrowing in clay soil) requires ~8x energy
compared to burrowing in sandy soil
!
Energy depends on soil type:
○
Access to food
!
Protection from predators
!
A more stable thermal environment
!
Burrowing required 360-3400x energy costs than walking, BUT supplies:
○
Burrowing
•
Energy required is 3.4 x BMR
!
Ex. Herring gull, wandering albatross
!
Gliding flight:
○
Large wings compared to body weight -extremely aerial species
!
Energy required is 3.4-6.4 x BMR
!
Ex. Purple martin, house martin, barn swallow, bank swallow, sooty
tern
!
Very active flight:
○
Without the large wings of extremely aerial species
!
Energy required is up to 14 x BMR
!
Active Flight:
○
Ex. European robin: ~23 x BMR
!
For very short flights (few seconds), energy requirements are even higher
○
Energy costs of flight in bats is very similar to that of birds
○
Ex. 10g bird flying 1km has <1% cost compared to 10g mouse
walking/running 1km
!
Energy costs are high/unit time but low/unit distance
○
Flying
•
Less energy required compared to running or flying (per unit distance) in
terrestrial mammals and birds
○
Most energy efficient speed (per unit distance) is ~0.5 m/s (for most
aquatic mammals and birds)
!
Surface swimming -energy required increases with speed
○
~0.6 m/s for 1kg animal
□
~1.3 m/s for 60kg animal
□
Most efficient speed:
!
Submerged swimming -more energy efficient at faster speeds (less
drag/resistance)
○
~2 x RMR (captive animals)
!
~6-9.3 x RMR in wild penguins catching prey
!
Energy required when swimming at most energy efficient speeds:
○
Energy for high speed swimming to catch prey is up to 50 x
RMR
□
Energy cost is too high --> starvation
□
Sea lion (45kg) with fishing net (0.6kg): energy swimming is 4.3x
higher with net
!
Energy costs of swimming are greatly increased when fishing net, plastic
buoys, or garbage become attached to marine mammal or bird
○
Swimming
•
Swinging from tree to tree using arms
○
Ex. Spider monkey -x1.5 normal walking
!
Energy cost (kcal/kg/km)
○
Spider monkey/slow loris energy use increases ~65% compared to
resting
!
Hanging without movement:
○
3x more energy than 20% required by most standing animals compared to
lying
○
Brachiation
•
Energy required to capture (predators), manipulate, chew and ingest food
○
Ruminants: 20-59% increase over standing
○
Birds: ~11% increase over perching
○
Feeding
•
Fighting (hierarchy and mating rights)
○
Playing
○
Grooming
○
Ruminating
○
Escaping from predators
○
Other Activities:
•
Maintenance Energy -Activity
Energy: Maintenance & Activity
Monday,+February+ 6,+2017 11:37+AM
Document Summary
Metabolizable energy (me) - intake at which body weight and composition remain constant, in a healthy, non-reproducing animal/bird living in its normal environment. Daily energy expenditure (dee) = bmr + activity + thermoregulation + food metabolism. It is not the "average" but the "variation" that determines whether animals reproduce, survive or die. Maintenance energy/ daily energy expenditure - measures of metabolism. *nutrient/food metabolism = heat increment of feeding (hi) = specific dynamic. Requires energy, ~20% greater than lying (in mammals) Exceptions: equids (horse family), passively lock legs to remain standing (elephants) Birds, estimated at 17 +/- 11 % greater than sitting. Painted quail, bobwhite quail, chukar, partride, guinea fowl, wild turkey, domestic chicken. Energy cost (kcal/kg/hr) for a given speed is higher for smaller animals. Energy cost of increasing speed of 1km/ hr is greater in smaller animals. Gait may change when the energy cost of increasing speed using the slower gait is greater than changing to a faster gait.
