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 2 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 3 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 4 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 Copyright James D Thomson 6 An exception to Bergmanns Rule?
National Geographic Copyright James D Thomson 7 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 Copyright James D Thomson 8 Maximum SA:V ratio? Chrysopelea gliding snake, Borneo; restricted to warm tropics National Geographic, Tim Laman
Copyright James D Thomson 9 Minimum SA:V ratio? Pika, Ochotona princeps: alpine tundra rabbit; restricted to cold habitats; note spherical shape, reduced ears (for a rabbit) National Geographic Copyright James D Thomson 10 Variation in appendages: arctic and desert hares Lepus arcticus (Art Wolfe) Lepus californicus (www.pestproducts.com) Copyright James D Thomson 11 What else matters? Insulation more important than size/shape
www.dinosaur-museum.org
mercury.bio.uaf.edu Copyright James D Thomson 12 Thick insulation: Half-sheared sheep National Geographic Copyright James D Thomson 13 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
Copyright James D Thomson 14 Thick insulation: Cross-section of seal Blubber Guts Musculature 42% 58% of cross- sectional area Copyright James D Thomson 15 Hair and feathers provide adjustable insulation Video clip, Clarks Nutcracker on a cold morning in Colorado Copyright James D Thomson 16 What else matters? Convective cooling enhanced by vascularization Lepus californicus (National Geographic) L. americanus (JDT) Copyright James D Thomson 17 Another way to provide well- vascularized surface area for cooling Copyright James D Thomson 18 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 Copyright James D Thomson 19 What else matters? Convection enhanced by evaporation /www.bluegina.com Copyright James D Thomson 20 What else matters? Convection enhanced by evaporation /www.bluegina.com Copyright James D Thomson 21 Extra slide if time: Range dependence on other organisms: saguaro, gila woodpecker, cactus wren, elf owl www.southwestbirders.com Copyright James D Thomson 22 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 23 Copyright James D Thomson 24 Henry VIII: Low SA / V Mustela erminea: High SA / V Holbein Copyright James D Thomson 25 Henry VIII: Better? Mustela erminea: Worse? Copyright James D Thomson 26 Ecology 53(5): 939-943 (1972) Weasel Woodrat
Warm ((Cold)) Curls into flat disk Curls into ball Copyright James D Thomson 27 So, the weasel body plan is well-suited for warm climates! M.erminea !making their actual distribution paradoxical! Copyright James D Thomson 28 Mustela frenata and Thomomys talpoides, Rocky Mountain Biological Lab, D. Inouye photos Copyright James D Thomson 29 Typical weasel prey:
So, the paradox involves the requirements of the weasels predatory lifestyle
Copyright James D Thomson 30 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 31 Copyright James D Thomson