Lecture 20, Regulating the Internal Environment: I. Thermoregulation • • • • • Heat Gain or Loss is controlled by: Conduction: direct transfer of heat between molecules of the environment and those of body surface Convection:transfer of heat by movement of air or water past body surface Radiation: emission of electromagnetic radiation Evaporation: loss of heat from the surface of a liquid that is losing some of its molecules as gas • Ectotherms: (poikilotherms) derive heat mainly from their surroundings • Endotherms: (homeotherms) derive heat mainly from their metabolism Endothermy: Advantages & Disadvantages • Advantages: – Terrestrial environments are characterized by large temperature fluctuations – Endothermy liberates an animal from dependence on ambient temperature • Disadvantages: – Energetically very expensive: • Resting Homo sapiens: at 200C, consumes 1300 - 1800 kcal per day • Resting alligator of equivalent body mass, consumes 60 kcal per day 1 Thermoregulation: Mechanisms 1 1. Adjusting Rate of Heat Exchange Between an Animal & Its Environment • Body insulation: hair, feathers, fat • Vasodilation: enhances transfer of heat to environment • Vasoconstriction: reduces transfer of heat to environment • Counter-current heat exchanger Thermoregulation: Mechanisms 2 2. Cooling by Evaporative Heat Loss • Phase change from liquid water to water vapor releases a large amount of heat • Very effective when relative humidity is low to moderate • Mammals: sweat or pant • Birds: gular fluttering • Behavioral adaptations: – e.g., Bathing ----> Thermoregulation: Mechanisms 3 3. Behavioral Responses: • Particularly important in ectotherms 2 Endothermic Invertebrates • Pre-flight warm-up in large moths, beetles and bees • Contract all flight muscles in synchrony in order to heat up • Make use of countercurrent heat exchanger to maintain high thoracic temperature Endothermic Fish Endothermic Fish: Rete mirable (e.g., Bluefin tuna, swordfish, great white shark) 3 Torpor • State of reduced metabolic activity in which the heart rate and respiratory system slow down • Used by many endotherms as a mechanism to reduce energy consumption • Short-term torpor: exhibited on daily basis in small mammals (e.g., shrews) & birds (e.g., hummingbirds) • Long-term torpor: – Hibernation: response to cold (triggered by photoperiod) – Estivation: response to heat or drought “Ectothermic” Birds “Ectothermic” Mammals: Hibernation 4 Feedback Mechanisms in Thermoregulation Heat-shock Proteins • High body temperature can be particularly dangerous because many biologicallyimportant proteins begin to denature at ~420C. • In response to high temps, cells release heat shock proteins which help buffer other proteins against being denatured • Lee Weber, UNR, expert on heat shock proteins: – using cut-throat trout heat shock titers as an index of environmental quality Regulating Internal Environment II: Water Balance & Waste Disposal • Water balance & excretion are controlled by transport epithelia: a layer or layers of of specialized epithelial cells that regulate solute movements • Transport epithelia: – Either face the external environment – Or, a tubular channel that leads to the outside through an opening on the body surface – Usually arranged in tubular networks with extensive surface area 5 Waste Products of Metabolism Nitrogenous Wastes • Ammonia, the direct product of metabolism (via deamination of proteins & nucleic acids), is very toxic stuff (good for cleaning windows, however) • Ammonia, however, is very soluble in H2 O and passes easily through biological membranes • Because of its toxicity, ammonia can only be tolerated in very low concentrations Form of Nitrogenous Waste Disposal • Form of nitrogenous waste is associated with an animal’s phylogeny and habitat • Ammonia: restricted to aquatic organisms (FW fish, tadpoles); Requires lots of H2O • Urea: Mammals, adult amphibians, marine fish & marine turtles. 100,000x less toxic than NH3 • Uric acid: Birds, reptiles, land snails, insects. Relatively non-toxic and very insoluble in H2O; excreted as a paste. 6 Urea vs. Uric Acid: Importance of Reproductive Mode • Urea: Excreted in H2O as urine. In adult amphibians & mammals, produced by liver in reaction of NH3 & CO2. Energetically expensive but conserves H2O by reducing toxicity. • Uric Acid: Excreted as paste. Energetically expensive but very effective at conserving H2O. • What about embryos? – Amphibian eggs: urea (urine) can diffuse out of shell-less egg; – Mammalian fetus: urea can diffuse out into mother’s bloodstream; – Bird & insect eggs: shelled eggs are impermeable to liquids (urine); however, uric acid precipitates out of solution and stored in egg as a solid Terminology of Osmoregulation Marine fish: hypotonic osmoregulator: constantly loses water through osmosis and therefore has to drink constantly; Gills actively transport Cl- out 7 Freshwater Fish: hypertonic osmoregulator: constantly gains water through osmosis; produces copious quantities of dilute urine; gills actively transport Cl- in Osmotic Balance on Land • Desiccation: a major threat in terrestrial environments • Major sources of H2O loss: – Gas exchange organs – Urine • Mammals very greatly in their ability to minimize water loss via urine • Kangaroo rat: recovers 90% of water loss through metabolic H2O. • KR: 22.5% lost in urine • Human: 60% lost in urine Waste Products of Metabolism 8
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