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Chapter 5. Energy Expenditure and Fatigue. Measuring Energy Expenditure: Direct Calorimetry. Substrate metabolism efficiency 40% of substrate energy ATP 60% of substrate energy heat Heat production increases with energy production Can be measured in a calorimeter
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Chapter 5 • Energy Expenditure and Fatigue
Measuring Energy Expenditure:Direct Calorimetry • Substrate metabolism efficiency • 40% of substrate energy ATP • 60% of substrate energy heat • Heat production increases with energy production • Can be measured in a calorimeter • Water flows through walls • Body temperature increases water temperature
Measuring Energy Expenditure:Respiratory Exchange Ratio • O2 usage during metabolism depends on type of fuel being oxidized • More carbon atoms in molecule = more O2 needed • Glucose (C6H12O6) < palmitic acid (C16H32O2) • Respiratory exchange ratio (RER) • Ratio between rates of CO2 production, O2 usage • RER = VCO2/VO2
Measuring Energy Expenditure: Respiratory Exchange Ratio • RER for 1 molecule glucose = 1.0 • 6 O2 + C6H12O6 6 CO2 + 6 H2O + 32 ATP • RER = VCO2/VO2 = 6 CO2/6 O2 = 1.0 • RER for 1 molecule palmitic acid = 0.70 • 23 O2 + C16H32O2 16 CO2 + 16 H2O + 129 ATP • RER = VCO2/VO2 = 16 CO2/23 O2 = 0.70 • Predicts substrate use, kilocalories/O2efficiency
Energy Expenditure at Rest and During Exercise • Metabolic rate: rate of energy use by body • Based on whole-body O2 consumption and corresponding caloric equivalent • At rest, RER ~0.80, VO2 ~0.3 L/min • At rest, metabolic rate ~2,000 kcal/day
Energy Expenditure DuringMaximal Aerobic Exercise • VO2max expressed in L/min • Easy standard units • Suitable for non-weight-bearing activities • VO2max normalized for body weight • ml O2 kg-1 min-1 • More accurate comparison for different body sizes • Untrained young men: 44 to 50 versus untrained young women: 38 to 42 • Sex difference due to women’s lower FFM and hemoglobin
Anaerobic Energy Expenditure:Postexercise O2 Consumption • O2 demand > O2 consumed in early exercise • Body incurs O2 deficit • O2 required − O2 consumed • Occurs when anaerobic pathways used for ATP production • O2 consumed > O2 demand in early recovery • Excess postexercise O2 consumption (EPOC) • Replenishes ATP/PCr stores, converts lactate to glycogen, replenishes hemo/myoglobin, clears CO2
Anaerobic Energy Expenditure:Lactate Threshold • Lactate threshold: point at which blood lactate accumulation markedly • Lactate production rate > lactate clearance rate • Interaction of aerobic and anaerobic systems • Good indicator of potential for endurance exercise • Usually expressed as percentage of VO2max
Anaerobic Energy Expenditure:Lactate Threshold • Lactate accumulation fatigue • Ability to exercise hard without accumulating lactate beneficial to athletic performance • Higher lactate threshold = higher sustained exercise intensity = better endurance performance • For two athletes with same VO2max, higher lactate threshold predicts better performance
Measuring Anaerobic Capacity • No clear, V̇O2max-like method for measuring anaerobic capacity • Imperfect but accepted methods • Maximal accumulated O2 deficit • Wingate anaerobic test • Critical power test
Energy Expenditure During Exercise:Economy of Effort • As athletes become more skilled, use less energy for given pace • Independent of VO2max • Body learns energy economy with practice • Multifactorial phenomenon • Economy with distance of race • Practice better economy of movement (form) • Varies with type of exercise (running vs. swimming)
Energy Expenditure:Successful Endurance Athletes 1. High VO2max 2. High lactate threshold (as % VO2max) 3. High economy of effort 4. High percentage of type I muscle fibers
Fatigue and Its Causes • Fatigue: two definitions • Decrements in muscular performance with continued effort, accompanied by sensations of tiredness • Inability to maintain required power output to continue muscular work at given intensity • Reversible by rest
Fatigue and Its Causes • Complex phenomenon • Type, intensity of exercise • Muscle fiber type • Training status, diet • Four major causes (synergistic?) • Inadequate energy delivery/metabolism • Accumulation of metabolic by-products • Failure of muscle contractile mechanism • Altered neural control of muscle contraction
Fatigue and Its Causes:Energy Systems—PCr Depletion • PCr depletion coincides with fatigue • PCr used for short-term, high-intensity effort • PCr depletes more quickly than total ATP • Pi accumulation may be potential cause • Pacing helps defer PCr depletion
Fatigue and Its Causes:Energy Systems—Glycogen Depletion • Glycogen reserves limited and deplete quickly • Depletion correlated with fatigue • Related to total glycogen depletion • Unrelated to rate of glycogen depletion • Depletes more quickly with high intensity • Depletes more quickly during first few minutes of exercise versus later stages
Fatigue and Its Causes:Energy Systems—Glycogen Depletion • Fiber type and recruitment patterns • Fibers recruited first or most frequently deplete fastest • Type I fibers depleted after moderate endurance exercise • Recruitment depends on exercise intensity • Type I fibers recruit first (light/moderate intensity) • Type IIa fibers recruit next (moderate/high intensity) • Type IIx fibers recruit last (maximal intensity)
Fatigue and Its Causes:Energy Systems—Glycogen Depletion • Depletion in different muscle groups • Activity-specific muscles deplete fastest • Recruited earliest and longest for given task • Depletion and blood glucose • Muscle glycogen insufficient for prolonged exercise • Liver glycogen glucose into blood • As muscle glycogen , liver glycogenolysis • Muscle glycogen depletion + hypoglycemia = fatigue
Fatigue and Its Causes:Energy Systems—Glycogen Depletion • Certain rate of muscle glycogenolysis required to maintain • NADH production in Krebs cycle • Electron transport chain activity • No glycogen = inhibited substrate oxidation • With glycogen depletion, FFA metabolism • But FFA oxidation too slow, may be unable to supply sufficient ATP for given intensity
Fatigue and Its Causes:Metabolic By-Products • Pi: From rapid breakdown of PCr, ATP • Heat: Retained by body, core temperature • Lactic acid: Product of anaerobic glycolysis • H+ Lactic acid lactate + H+
Fatigue and Its Causes:Metabolic By-Products • Heat alters metabolic rate – Rate of carbohydrate utilization • Hastens glycogen depletion • High muscle temperature may impair muscle function • Time to fatigue changes with ambient temperature • 11°C: time to exhaustion longest • 31°C: time to exhaustion shortest • Muscle precooling prolongs exercise
Fatigue and Its Causes:Metabolic By-Products • Lactic acid accumulates during brief, high-intensity exercise • If not cleared immediately, converts to lactate + H+ • H+ accumulation causes muscle pH (acidosis) • Buffers help muscle pH but not enough • Buffers minimize drop in pH (7.1 to 6.5, not to 1.5) • Cells therefore survive but don’t function well • pH <6.9 inhibits glycolytic enzymes, ATP synthesis • pH = 6.4 prevents further glycogen breakdown
Fatigue and Its Causes:Lactic Acid Not All Bad • May be beneficial during exercise • Accumulation can bring on fatigue • But if production = clearance, not fatiguing • Serves as source of fuel • Directly oxidized by type I fiber mitochondria • Shuttled from type II fibers to type I for oxidation • Converted to glucose via gluconeogenesis (liver)
Fatigue and Its Causes:Neural Transmission • Failure may occur at neuromuscular junction, preventing muscle activation • Possible causes – ACh synthesis and release • Altered ACh breakdown in synapse • Increase in muscle fiber stimulus threshold • Altered muscle resting membrane potential • Fatigue may inhibit Ca2+ release from SR
Fatigue and Its Causes:Central Nervous System • CNS undoubtedly plays role in fatigue but not fully understood yet • Fiber recruitment has conscious aspect • Stress of exhaustive exercise may be too much • Subconscious or conscious unwillingness to endure more pain • Discomfort of fatigue = warning sign • Elite athletes learn proper pacing, tolerate fatigue