BA

Module 2

Energy is the capacity to work.

Work is the transfer of energy by a force acting on an object as it is displaced.

  • work = force x distance

  • force = mass (kg) x acceleration

  • acceleration = (velocity)²

  • velocity = distance (m) / time (s)

  • both work and energy are measured: J = kg x m² / s²

Power is the rate at which work is done

  • W = J/s

Metabolism is the set of processes by which cells and organisms acquire, rearrange, and void commodities in ways that sustain life.

Metabolic rate is an animal’s rate of energy consumption; the rate at which it converts chemical-bond energy to heat and work—a measure of power.

2 main energy storage forms:

  • reducing energy (NADH and FADH2)

  • high energy bonds (ATP)

High energy bonds:

  • energy released when covalent bonds are broken

  • ATP and GTP

Carbohydrates:

  • many hydroxyl groups (OH-)

  • glucose is the most common carbohydrate in animal diets

Monosaccharides:

  • used for energy and biosynthesis

  • have three to seven carbons

Polysaccharides:

  • long chain of monosaccharides

  • used for energy storage (glycogen) and structural molecules (chitin)

Glycogen is the main carbohydrate storage form in animals (vertebrates).

Glycogen synthesis is glycogenesis; glycogen breakdown is glycogenolysis.

  • reciprocal regulatory enzymes act on glycogen synthase or glycogen phosphorylase

  • when synthesis is turned on, phosphorylase is turned off

Glycolysis:

  • takes place in the cytoplasm

  • anaerobic

  • produces reducing equivalents and ATP

  • end produce is pyruvate

Lactate and amino acids can also be converted to pyruvate. Pyruvate is carried into the mitochondria.

Pyruvate dehydrogenase (PDH) oxidizes pyruvate to form acetyl coA and NADH in the mitochondria.

Oxidation of NADH must occur for glycolysis to proceed.

In the presence of O2, NADH is oxidized to NAD+

  • alpha-glycerophosphate shuttle in invertebrates

  • malate-aspartate shuttle in vertebrates

  • from cytoplasm to mitochondria

In the absence of O2, NADH cannot be used rapidly

  • build up in NADH means a drop in NAD+, inhibiting glycolysis

  • lactate dehydrogenase (LDH) converts pyruvate, NADH, and H+ to lactate and NAD+

  • reversible process

  • other pathways form less toxic end products and more ATP than lactate

    • ex. succinate and proprionate in invertebrates

Lipids:

  • hydrophobic

  • carbon backbone in linear or ring structure

  • used for energy metabolism, cell structure, and signaling

Fatty acids:

  • chain of carbons with a carboxyl end

  • saturated: no double bonds

  • unsaturated: one or more double bonds

Beta-oxidation:

  • fatty acids are dense, more reduced forms of carbon (require more oxidation) to unlock energy

  • takes place in the mitochondria, aerobic metabolism

  • results in the formation of Acetyl CoA

  • Acetyl CoA is then oxidized

Ketones:

  • tissues that cannot metabolize fatty acids metabolize ketones

    • ex. brain and shark muscle

  • ketogenesis:

    • fatty acids converted to Acetyl CoA

    • Acetyl CoA converted to ketones

    • ketone bodies move through circulation

  • ketolysis:

    • ketones are broken down into Acetyl CoA

    • Acetyl CoA then participates in oxidative phosphorylation

Mitochondrial Metabolism

  • glucose → metabolites

  • metabolites → Acetyl CoA

  • Acetyl CoA enters the krebs cycle and is oxidized to form reducing agents

  • reducing agents are oxidized to release energy

  • O2 is the final electron acceptor

Parasites lack mitochondrial genome and loss their ability to rely on O2.

Tricarboxylic Acid (TCA) Cycle:

  • generates reducing equivalents

  • amphibolic pathway: some intermediates are used for synthesis while others are used for breakdown

Electron Transport System (ETS):

  • within the inner mitochondrial membrane

  • composed of four multisubunit proteins (complexes I, II, III, IV)

  • composed of two electron carriers (ubiquinone and cytochrome c)

  • oxidation: 4e- + 4H+ + O2 → 2H2O

  • generates a proton gradient, heat, water, and ROS

ATP Synthesis:

  • ADP + P → ATP

  • proton motive force

  • F1F0ATPase uses energy in proton force to produce ATP

  • no physical link

  • functionally coupled

Phosphocreatine:

  • alternative high-energy compound

  • creatine + ATP ←→ ADP + phosphocreatine

  • phosphocreatine can move throughout the cell like ATP

  • enflux of high energy

Spectroscopy:

  • measures ATP turnover

  • pros:

    • measures cellular energy currency

    • accurate over short time scales

  • cons:

    • difficult

    • subject must be restrained

    • equipment not portable

Hess’ Law

  • total amount of energy released for breakdown of given amount of fuel always the same

  • regardless of intermediate chemical steps

Direct Calorimetry:

  • measurement of heat during chemical/physiological processes

  • pros:

    • quite accurate

    • accounts for aerobic and anaerobic energy production

  • cons:

    • restrained

    • heavy and complicated

    • makes assumptions about anabolic vs catabolic actvitity

Indirect Calorimetry: Respirometry

  • inferring metabolic heat production

  • pros:

    • friendly and easy

    • can be portable

    • very accurate with assumptions

    • easy on active animals

  • cons:

    • aerobic metabolism only

    • must sample gases

    • animal must be ‘tied’ at least

Respirometry Quotient:

  • rate of CO2 production/O2 consumption

  • RQ for carbohydrates > proteins > lipids

  • differ depending on which fuel is being oxidized

  • a greater RQ occurs during anaerobic exercise

  • fats produce less CO2 per O2 consumed

  • 14.9-18.7% more O2 required when oxidizing fats compared to carbs

Hummingbirds hovering VO2 will be 15-19% greater when fasted (burning fat) compared to when fed (burning carbs). When fasted, they are burning fats which require more oxygen to burn.

Chamber Respirometry:

  • animal enclosed in chamber

  • pros:

    • easiest approach

    • more accurate

    • easier to control

  • cons:

    • contained animal

    • risk of asphyxiation

    • can be messy (all functions in the chamber)

  • closed system:

    • make sure all gases are accounted for

    • more accurate with longer time scales

    • switching between rest and activity complicated the study

Mask Respirometry:

  • only covers mouth/nose/head

  • small volume, faster flow rates, animal can behaviour nearly naturally

  • poorer signal to nose ratio, composition of gases harder to control, assumes gas concentrations

The ATP turnover rate and the rate of fuel delivery to or supply in the cell do not necessarily match instantly. ATP is used instantly, while fuel delivery and processing take time.

O2 consumption/CO2 production measured at the respiratory interface with the environment does not reflect instantaneous O2 consumption/CO2 production in tissues.

  • during transition between exercise and rest, there can be an offset in tissue level O2 consumption or metabolic rate, and VO2

  • actual rate of oxygen consumption and CO₂ production in tissues is not reflected instantly in the air you breathe

Inverse relationship between max power and sustainability.

  • at maximum power, energy is used instantly

  • ATP Pool → Phosphocreatine → Anaerobic Glycolysis → Aerobic Glycolysis → Aerobic Lipolysis

O2 consumption remains high even after returning to low activity levels as glycogen stores are replenished.

  • Lactate produced is used to…

    • Cori Cycle: Gluconeogenesis in the liver to return glucose to liver and rebuild glycogen

    • Lactate Shuttle: Used as fuel → pyruvate → Acetyl CoA

  • Creatine produced is used to…

    • rebuild phosphocreatine at the expense of ATP

    • since ATP is largely being generated using aerobic metabolism, this can be done

Diving:

  • O2 consumption may not be possible

  • reduce exchange of O2/CO2

  • hypoxic (low oxygen) and hypercapnia (high carbon dioxide)

  • perform compensatory hyperventilation → store a lot of O2 in tissues and “blow off” CO2

Basal Metabolic Rate (BMR):

  • metabolic rate of homeothermic animal at rest

  • at a temperature within thermoneutral zone

Standard Metabolic Rate (SMR):

  • BMR for poikilothermic animal at a defined environmental temperature

Resting Metabolic Rate (RMR):

  • metabolic rate when animal is at rest under defined conditions

  • slightly higher than BMR

Maximum Aerobic Metabolic Rate (VO2 max):

  • maximum sustainable VO2

  • during intense exercise or when a homeotherm is exposed to very cold temperatures

Supermaximal Metabolic Rate:

  • burst only

Field Metabolic Rate:

  • the actual realized metabolic rate when the animal is behaving naturally in the wild

Daily Energy Expenditure:

  • total energetic cost of a day of life

  • not a rate, just an energy amount

Allometric scaling describes metabolic rate, where one variable scales to the power of the other.

  • whole-animal metabolic rate increases with increasing size (larger animals have more cells)

  • mass-specific metabolic rate decreases with increasing size (smaller animals have a larger surface area to mass ratio which means they burn more energy per unit of body mass)

  • the exponent for one is the inverse of the other

  • Scaling Exponent (b) tells you how a trait changes with body size. It’s a power that shows whether the trait increases faster, slower, or proportionally with size. About 0.67 for mammals and 0.75 interspecifically.

  • Scaling Coefficient (a) is a constant that sets the starting point for the scaling. It determines how large or small the value of the trait is to begin with.

Surface area to volume scaling affect gas exchange and nutrient absorption too.

VO2 max scales at an exponent greater than BMR but it is limited by O2 absorption and distribution.