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CHAPTER 42 CONTROL OF ENERGY BALANCE, METABOLIC RATE, AND BODY
TEMPERATURE Prepared by Brenda Leady, University of Toledo Copyright (c) The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
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All animals require sources of carbon, nitrogen, and other nutrients
Used to assemble macromolecules or generate ATP Animals must have mechanisms to maintain blood levels of important nutrients despite irregular nutrient flow
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Metabolism – all the activities and chemical reactions in an animal’s body
Metabolic rate – rate at which animal uses fuel to supply ATP for these reactions Metabolism and body temperature are related – heat is generated as a by-product of nutrient breakdown and ATP synthesis
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Nutrient utilization and storage
Nutrient utilization divided into 2 alternating phases Absorptive phase – occurs when ingested nutrients enter blood stream from GI tract Some of ingested nutrients used for immediate energy needs, others stored Postabsorptive stage – occurs when GI tract empty of nutrients so body’s stores used
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Absorbed carbohydrates
Chief monomers absorbed are glucose, galactose, and fructose Glucose is 1 of 2 major energy sources during absorptive phase (other is fat) Much of absorbed glucose used to synthesize ATP Skeletal muscle major consumer of glucose Can store as glycogen Excess glucose stored as glycogen in liver or triglycerides
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Triglycerides Too large to diffuse across plasma membrane of intestinal epithelial cells Digested into monoglycerides and fatty acids Diffuse into intestinal epithelial cells Resynthesized into triglycerides Packaged into chylomicrons for blood transport Lipoprotein lipase releases fatty acids to diffuse into cells of the body Some ingested fat used during absorptive phase for energy
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Amino acids Taken up by all body cells Used to synthesize proteins
Excess amino acids not stored as protein Excess converted by liver cells into carbohydrate or fat
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Postabsorptive period
Synthesis of glycogen and fat slows and breakdown begins Blood glucose concentration maintained Reactions that provide glucose to blood Using fat instead of glucose for energy (glucose sparing)
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Reactions that provide glucose to blood
Glycogenolysis – glycogen hydrolysis in liver Gluconeogenesis – liver converts noncarbohydrates into glucose Lipolysis releases fatty acids (alternate energy source to glucose) and glycerol (converted to glucose by liver) Proteins can also be converted to glucose in prolonged fasting
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Glucose sparing Most organs and tissues increase fat utilization for energy Reserves glucose for use by nervous system Essential step is lipolysis Fatty acids used by cells for energy Liver uses fatty acids to make ketones Prolonged energy source for many tissues, including brain Less protein breakdown required
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Regulation of phases Insulin Made by pancreas
Binds to cell surface receptor, stimulates signaling pathway, facilitates glucose diffusion into cells Glucose transporters (GLUTs) in intracellular vesicles move to the surface Also inhibits glycogenolysis and gluconeogenesis in the liver
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GLUT Proteins Transport Glucose in Animal Cells
All animal cells require transporters to move glucose across plasma membranes GLUTs make up a family of at least 14 related proteins Expressed in different tissues Structures similar across phyla – gene duplication of common ancestral gene Different GLUTS vary in ability to bind glucose High affinity – binds even at low glucose concentrations Muscle and fat have low affinity GLUT4 (also only one requiring insulin) Brain cells have high affinity GLUT
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Controlling insulin secretion
Blood glucose rises during absorptive state so insulin secretion rises Blood glucose levels fall during postabsorptive state so insulin secretion falls Negative feedback Input from nerve cells also plays role Eating meal increases insulin secretion Stressful situation decreases insulin secretion
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Maintaining blood glucose levels
Blood glucose must remain within the normal homeostatic range Too low and even high affinity GLUT receptors in the brain will not take in glucose Glucagon – from the pancreas stimulates glycogenolysis, gluconeogenesis, and ketone synthesis
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Energy expenditure Amount of fuel used during a given period of time to power all of its metabolic requirement Breakdown of organic molecules liberates energy to be transferred to ATP Not all energy used for ATP to do work Some energy appears as heat
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Metabolic rate Calorie – amount of heat required to raise the temperature of 1 gram of water 1 degree Celsius Commonly measured in kilocalories (kcal) Metabolic rate – total energy expenditure of an animal per unit time
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Basal metabolic rate (BMR) – at rest, in postabsorptive state, and at a standard temperature
Endotherms – generate their own heat and maintain relatively constant body temperature – BMR measured in thermoneutral zone Ectotherms – body temperature changes with environment – BMR measured at standard temperature for each species SMR or standard metabolic rate
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BMR estimation Direct calorimetry Indirect calorimetry
Energy expenditure and heat production directly related Measures heat given off by animal Indirect calorimetry Measures rate of oxygen use by animal Underestimates BMR due to anaerobic metabolism
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Not all tissues use oxygen or produce heat at the same rate
Metabolic rates of different tissues can vary independently of each other
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Factors impacting metabolic rate
Altered skeletal muscle activity Food-induced thermogenesis – greatest effect from ingested protein increasing heat production Mass-specific BMR – amount of energy expended per gram of body mass – exponential curve Smaller animals lose heat more rapidly so higher BMR Does not explain the same relationship in ectotherms
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Smallest endotherms face the special challenge of fueling their very high mass-specific metabolic rates Torpor – nightly lowering internal body temperature to just a few degrees above that of the environment Hibernation – extended for months
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Balance Body weight remains stable when amount of energy consumed equals the energy expended Appetite and metabolism change when food intake is more or less than the amount needed to maintain the body’s set point
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Food intake Satiety (fullness) – involved in short term control of feeding Satiety signals – remove sensation of hunger and set time period before hunger returns again Stretch receptors in the stomach and small intestines send signals to the brain Stomach and small intestine release hormones to suppress appetite
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Leptin – produced in adipose tissue in proportion to fat mass
Long-term control of food intake mediated by many different brain molecules, by hormones, and by emotional state, particularly in humans Leptin – produced in adipose tissue in proportion to fat mass As more fat is stored in the body, more leptin is secreted into the blood Acts on brain centers to reduce appetite and increase metabolic rate Decreased fat mass decreases leptin which decreases BMR and increases appetite
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Coleman Revealed a Satiety Factor in Mammals
Parabiosis – surgical connection between bloodstreams of 2 animals 1st experiment – connected wild type mouse to ob mouse Ob mouse ate less and gained less weight than usual Blood of wild type mouse contained factor signaling sufficient fat stores 2nd experiment – connected wild type mouse to db mouse Db mouse continued to gain weight while wild type mouse lost weight Obesity phenotype the same – factor was absent in ob mice and present but unable to act in db mice Most obese humans like db mice – produce leptin but fail to respond to it
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Regulation of body temperature
Most animals can only survive a narrow range of temperatures Chemical reactions depend on temperature Enzymes have optimal temperature Proteins can become denatured at high temperatures Heat alters structures of plasma and intracellular membranes
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Extreme cold is better tolerated
Ice crystals can rupture cell membranes Some animals can block ice crystal formation in their cells
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Maintaining body temperature
Classify animals according to Source of heat – internal or environmental Ability to maintain body temperature – fluctuating (heterotherms) or stable (homeotherms) Endothermic homeotherms – birds and mammals Ectothermic heterotherms – other vertebrates and most invertebrates Not all animals can be classified neatly all the time Hibernating animal (heterothermic) or deep ocean fish (homeothermic)
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Birds and mammals can quickly adjust the body’s mechanisms for retaining or releasing heat so that body temperature remains stable A suddenly awakened mammal is instantly capable of intense activity A reptile must wait until it can warm itself using the environment
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Disadvantages of endothermy
Must consume larger amounts of food to produce sufficient heat by metabolic processes Run the risk of overheating during intense activity, even in cold weather
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Heat exchange Radiation Conduction Convection Evaporation
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Radiation Emission of electromagnetic waves by the surfaces of objects Rate determined by temperature of radiating surface Conduction Body surface loses or gains heat through direct contact with cooler or warmer substances Water has a higher heat capacity than air Surface area important – elephant ears or bat wings
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Convection Evaporation
Transfer of heat by the movement of air or water next to the body Sitting near fans, bat flapping wings, or elephant waving ears Evaporation Water vaporizing from surface conducts heat used to drive process away from surface
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Heat gain or loss Core temperature of endotherms remains constant but skin surface varies considerably Blood flow to the skin can increase or decrease heat loss
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Countercurrent heat exchange
Conserves heat Dolphin flipper or bird legs Heat moves from warm arteries to adjacent veins carrying cooler blood Reduces heat lost to environment Fish keep muscles warm without losing heat through gills
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Behavioral adaptations
Evaporative heat loss Change rate of evaporation via perspiration Humidity determines rate of actual evaporation Panting, licking skin, bathing in water are alternatives to sweat glands Behavioral adaptations Changing exposed surface area – curling into ball, huddling, hunching Changing surroundings – critical for endotherms obtaining or losing heat
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Increased heat production
Changes in muscle activity major control in endotherms Shivering thermogenesis – skeletal muscle contractions without locomotion Nonshivering thermogenesis – primarily in brown fat’s mitcohondria where uncoupling proteins use H+ gradient to generate heat, not ATP synthesis
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Impact on public health
Obesity Body Mass Index (BMI) of 30 or greater Lack of physical activity reduces daily energy expenditure “Apples” versus “pears” – location of fat has health consequences Genetic factors – thrifty genes may have been survival mechanism Underweight Anorexia nervosa – reduce food intake to the point of starving, biological and psychological causes Bulimia nervosa – recurrent episodes of fasting and overeating, may include self induced vomiting or laxative use Treatments rely on counseling, nutritional education and medication
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