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Metabolism. Chapters 5-7. Metabolism. Sum of all chemical reactions occurring in a living organism Energy Conversion Chemical energy → (Electrical or Mechanical Energy) → Heat Material Conversion Catabolism - break down of complex substances Anabolism - build up of complex substances.
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Metabolism Chapters 5-7
Metabolism • Sum of all chemical reactions occurring in a living organism • Energy Conversion Chemical energy → (Electrical or Mechanical Energy) → Heat • Material Conversion • Catabolism - break down of complex substances • Anabolism - build up of complex substances
Uses of Energy • Biosynthesis • Replacing body structures • Growth • Reproduction • Storage (Fat, Glycogen) • Exported Materials • Maintenance (homeostasis) • External Work (e.g, movement)
Energy Metabolism • Law of Entropy(2nd Law of Thermodynamics) • all metabolic processes involve a loss of free energy (organized energy disorganized energy) • Animals require a constant input of organized energy (organic chemical bonds) • All energy involved in metabolism is eventually lost in the form of heat Fig 5.1
Energy Metabolism • Energy usage by an organism • Rate at which organized energy is converted into heat • Calculation of Metabolic Rate • Direct calorimetry • measure heat production (kJ or Cal) • Indirect calorimetry • measure chemical changes • C6H12O6 + 6O2 6CO2 + 6H2O + Energy (673 Cal, 2820 kJ) Fig 5.4, Box 5.4
Indirect Calorimetry:Oxygen Consumption • Almost synonymous w/ metabolism • NOTE: only indicates energy usage through aerobic respiration • Accurate measure of energy expenditure through aerobic respiration • Roughly equal heat generation per liter O2 by carbohydrates, fats and proteins Table 5.1
Indirect Calorimetry: Carbon Dioxide Production • Amount of CO2 formed does not always equal amount of O2 consumed • Respiratory Quotient (RQ) • Amt CO2 produced/O2 consumed • Varies for different energy sources Table 5.2
O2 vs. CO2 • CO2 production is not as effective a measure of energy metabolism as O2 consumption • Energy yield per ml CO2 produced varies greatly • CO2 production can change easily through non-metabolic processes • e.g. hyperventilation
What Affects Metabolic Rate? • Physical Activity • Environmental Temperature • Digestive Processing (Specific Dynamic Action) • Body Size • Age • Gender • Endocrine Activity • Circadian Rhythms • Aquatic Salinity (Osmoregulation) Fig 5.5
Measuring “Apples and Apples” Metabolic Rate • Define physiological conditions under which metabolism is measured • Basal metabolic rate (BMR) – homeotherms • Temperature in thermal neutral zone • Fasting • Resting • Standard metabolic rate (SMR) – poikilotherms • Fasting • Resting
Metabolism and Body Size • Kleiber’s Rule • For eutherian mammals • Oxygen Consumption (VO2) = 0.676(Mass)0.75 • Specific Oxygen Consumption (VO2/kg) = 0.676(Mass)-0.25 • Small animals have relatively higher metabolic rates • E.g. shrews have 100x the per-gram VO2 as an elephant Figs 5.6-5.10
Metabolism and Body Size • Marsupial Mammals • VO2 = 0.409(Mass)0.75 • Passerine Birds • VO2 = 1.11(Mass)0.72 • Non-Passerine Birds • VO2 = 0.679(Mass)0.72 • Other Organisms • Ectothermic vertebrates • Invertebrates • Protozoa • Plants
Why Does Metabolic Rate Scale to Mass0.75? • Max Rubner – study on dogs • Small and large dogs have same body temperature • Heat must be produced in relation to heat loss • Heat production per square m2 surface area equal in small and large dogs • Large dogs have relatively lower surface areas • Rubner’s Surface Rule • Metabolic rate (heat production) surface area
What’s Wrong With This? • If metabolism was directly related to scaling of heat loss, it should scale to Mass0.67 • If related to heat generation and body temperature maintenance, why is it seen in ectothermic organisms?
Why Does Metabolic Rate Scale to Mass0.75? • O2 delivery mechanisms function Mass0.75 • Lung Ventilation Mass0.75 • Lung Volume Mass1.0 • Breathing Rate Mass-0.25 • Cardiac Output Mass0.75 • Heart Mass Mass1.0 • Heart Rate Mass-0.25 • Do these cause metabolism’s scaling, or does metabolism cause their scaling?
Why Does Metabolic Rate Scale to Mass0.75? • West et al. model- space filling fractal model • Biological distribution networks have a fractal design (branching) • Delivery of volumes of material to tissues approximated as spheres • Account for number of branchings needed to fill a given body volume (mass) , change in diameter, and delivery, flow to tissues α mass0.75 • Supply limitation Fig 5.12a
Why Does Metabolic Rate Scale to Mass0.75? • Darveau et al. model- allometric cascade • Overall MR = Σ various contributors to ATP turnover (materials supply and energetic demand) • b = Σ scaling exponents of these contributors • Scaling differs depending upon particular biochemical and physiological pathways activated • e.g., SMR - scaling dominated by demand (ATP usage) • e.g., MMR - both supply and demand influence scaling (O2 delivery vs. ATP usage) Fig 5.12b
Other Explanations • Related to the noncoding DNA content of cells (Koslowski et al. 2003) • Larger organisms have more noncoding DNA • More noncoding DNA produces larger cells • Larger cells have relatively lower MR • Mitochondrial function (Porter 2001) • Relatively less inner mitochondrial membrane surface in the cells of larger animals • Less ATP turnover • Less proton leak