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.