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Lecture 3: Physiological ecology of


animals: heat balance
1. Example of one environmental challenge to
physiology: Heat balance/thermal ecology of
animals
a. Modes of heat gain and loss; homeostasis
b. Size, shape, insulation, evaporative cooling
c. Tradeoff principle and adaptive compromises
(example of weasel body shape)

Copyright James D Thomson
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Radiationheat transfer by electromagnetic
radiation
Conductiondirect contact with substrate (e.g.,
feet lose heat to ground)
Convectionheat transfer mediated by moving
fluid (usually air or water)
Evaporationefficient cooling from wet surfaces
Redistributioncirculatory system redistributes
heat among body parts, esp. core to
appendages
One selected limiting factor:
heat budgets
Copyright James D Thomson
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Size matters to heat balance (and other
balances of gains and losses)
Homeostasis and surface area:volume ratio
Surface area determines equilibration rate
Volume provides the inertia
Radius: 1 2 3
SA/V: 3 1.5 1
Equilibrates most slowly
Equilibrates
quickly
Copyright James D Thomson
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Size matters
Bergmanns rule: Homeotherms tend to
be larger at higher latitudes (colder)

Helarctos malayanus
65 kg, short fur
Ursus americanus
275 kg, medium fur
Ursus maritimus
650 kg, long fur
All Wikipedia Copyright James D Thomson

Fossil record: horses shrank during two past
episodes of climate warming, grew during ice ages
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An exception to Bergmanns Rule?

National Geographic
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Shape matters:
Allens rule: extremities reduced in cold
climates
Sphere has least SA:V, so why shouldnt
homeotherms be spheres in cold climates?
Sometimes SA is needed for function
Sometimes particular shapes are needed
for function
Tradeoffs and adaptive compromises
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Maximum SA:V ratio?
Chrysopelea gliding snake, Borneo;
restricted to warm tropics
National Geographic, Tim Laman

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Minimum SA:V ratio?
Pika, Ochotona princeps: alpine tundra rabbit;
restricted to cold habitats; note spherical shape,
reduced ears (for a rabbit)
National Geographic
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Variation in appendages: arctic and
desert hares
Lepus arcticus (Art Wolfe) Lepus californicus (www.pestproducts.com)
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What else matters? Insulation more
important than size/shape

www.dinosaur-museum.org

mercury.bio.uaf.edu
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Thick insulation: Half-sheared sheep
National Geographic
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What else matters? Shaving! Can aquatic
mammals use fur for insulation? Drag is a drag.
When shaving is not enough:
Michael Phelps, 16 22 Olympic medals
HGH?
Anti-
drag
suit

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Thick insulation: Cross-section of seal
Blubber
Guts
Musculature
42%
58%
of cross-
sectional
area
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Hair and feathers provide
adjustable insulation
Video clip, Clarks Nutcracker on a cold
morning in Colorado
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What else matters? Convective
cooling enhanced by vascularization
Lepus californicus (National Geographic) L. americanus (JDT)
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Another way to provide well-
vascularized surface area for cooling
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What else matters? Countercurrent
circulation to limbs conserves heat
Arteries and veins should be appressed in appendages
to conserve heat; separated in appendages designed to
shed heat (Compare Ricklefs Fig. 3.24)
Countercurrent flow maintains gradient, so heat is
always flowing from outgoing blood to incoming blood
Outside (0!"
Body (35!"
30 20
5 10
15
Body (35!"
30 15
25 10
0
Vessels not appressed,
returning blood is chilled
Vessels appressed,
returning blood is warmer
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What else matters?
Convection enhanced by evaporation
/www.bluegina.com
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What else matters?
Convection enhanced by evaporation
/www.bluegina.com
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Extra slide if time: Range dependence on other organisms:
saguaro, gila woodpecker, cactus wren, elf owl
www.southwestbirders.com
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Reconciling an apparent paradox:
weasels are small predators, short-furred,
very long and thin
Copyright James D Thomson
Weasel in winter = ermine: active all year;
camouflaged for snowy environment
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Henry VIII: Low SA / V Mustela erminea: High SA / V
Holbein
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Henry VIII: Better? Mustela erminea: Worse?
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Ecology 53(5): 939-943 (1972)
Weasel Woodrat

Warm
((Cold)) Curls into flat disk
Curls into ball
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So, the weasel body plan is well-suited
for warm climates!
M.erminea
!making their actual distribution paradoxical!
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Mustela frenata and Thomomys talpoides, Rocky Mountain Biological Lab, D. Inouye photos
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Typical weasel prey:

Pocket gopher,
Thomomys talpoides

Seldom leaves
underground burrows;
mostly eats roots, tubers

So, the paradox involves
the requirements of the
weasels predatory
lifestyle

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Example of a tradeoff:
Being long and thin makes weasels subject to
thermal stresses (costly)!
!but allows them to be better predators
(beneficial).
Because they are long and thin, we infer that the
fitness gains of being a good hunter offset the
fitness costs of an expensive metabolism
Phenotypes of all organisms are riddled with
compromises dictated by tradeoffs

Copyright James D Thomson
Two reasons why natural selection
produces deeply imperfect organisms
Tradeoffs
Being good at x may necessarily imply being
bad at y
Constraints
Selection builds on what is already there,
especially existing developmental programs
Tinkering, yes; fundamentally fresh redesign,
no
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Copyright James D Thomson

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