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Transcript
Metabolism Continued Aerobic Respiration • Principle means of synthesizing ATP in animals • Three stages 1. Glycolysis (cytosol) • glucose pyruvate 2. Krebs Cycle (mitochondria) • formation of electron carriers and CO2 3. Oxidative Phosphorylation (mitochondria) • electron carriers create proton gradient used to generate ATP How Much ATP Can Be Generated? • 4 ATP gross (2 ATP net) in glycolysis • 2 GTP in the Krebs cycle • Theoretical maximum P/O ratios (#ATP per molecule of O2 consumed) of 3 ATP per NADH and 2 ATP per FADH2 – 10 NADH 3 = 30 ATP – 2 FADH2 2 = 4 ATP • Maximum yield = 38 ATP per glucose How Much ATP is REALLY Generated? • Less than 38 ATP (~30 in humans) • Most cells transfer electrons from cytosolic NADH to FADH2 in the mitochondrial matrix – Lose 2 ATP • Proton leakage across inner mitochondrial membrane – Lower actual P/O ratios: ~2.5 for NADH and 1.5 for FADH2 Other Ways of Generating ATP • Anaerobic Fermentation – – – – – Glycolysis used to generate ATP NAD+ reduced to NADH Must oxidize NADH back to NAD+ Reduce pyruvate into lactate Aquatic invertebrates • more complex pathways • Involve Krebs cycle reactions and truncated electron transport activity Anaerobic Metabolism • Problems – Low energy yield – Acid production affects cell/body pH • What do you do with it? – Reuse it • Lactate used by liver to regenerate glycogen (Cori cycle) – Get rid of it • Carp convert lactate to ethanol and release it through gills • Aquatic invertebrates release various organic molecules Other Ways of Generating ATP • Phosphagen Usage – Molecules store high energy phosphate groups • Arginine phosphate (invertebrates) • Creatine phosphate (vertebrates) – Transfer PO4 groups to ADP as ATP/ADP ratio lowers – Take up PO4 groups from ATP as ATP/ADP ratio increases Other Ways of Generating ATP • Stored Oxygen – Gas-binding pigments in tissues (e.g., myoglobin) can provide a reservoir of oxygen for aerobic respiration – Release O2 if intracellular PO2 drops Metabolism in Low O2 • Metabolism is independent of O2 concentrations to some degree (O2 regulation) • Low O2 may affect metabolism (O2 conformity) Responses to Low O2 • Increase ability to uptake O2 • Increased tolerance of hypoxia – Reliance on anaerobic metabolism Metabolism and Locomotion • Types of Locomotion – Cursorial – Swimming – Flight • How do these compare in energetic efficiency? Factors Influencing Cost of Locomotion • Support for body weight provided by the media – e.g. water – high support of body weight – e.g. air – low support for body weight • Resistance to movement – Dependent on density and viscosity of media – e.g. water – high resistance – e.g. air – lower resistance Cursorial Movement • Use limbs as levels to push against solid substrate • More energy required to run at higher velocities – Generally linear increase – Curvilinear at high speeds Cursorial Movement • Different patterns of limb movement (gaits) most efficient at different speeds • With increased speed, gait transitions occur – E.g. humans: walk run – E.g. horses: walk trot gallop Energetic Cost of Transport • COT = O2 consumed/distance traveled – Certain gaits are most efficient at a set speed – E.g. horses: run at speeds in each gait that minimize cost of transport Cost of Transport and Body Size • Small animals tend to have greater increases in energy expenditure with increasing velocity • Energetic cost of transport higher for smaller animals • Similar relationship among diverse animal taxa Flight • U-shaped relationship between O2 consumption and flight speed – O2 consumption minimized at a certain flight velocity – O2 consumption at higher AND lower speeds • Wing beats generate thrust and lift – Bernoulli effect – speed, lift – At low speeds, more lift has to be generated by downward beating of wings Cost of Flight • Speed of lowest cost of transport speed of minimum VO2 • e.g. parakeets – min VO2 at 35 kph – Min COT (VO2*kg-1*km-1) at 40 kph Swimming • Dense, viscous medium • Supports body mass • Generates high levels of drag – Force exerted in opposite direction of movement • Affected by media density, shape, size and velocity – – – – w/ density and viscosity with streamlining Drag surface area Drag velocity2 Which is the Most Efficient? • Swimming has the least expensive COT – Low speed, but no need for body support • Flight has the next least expensive COT – High energy input required, but high speeds and low drag increase efficiency