Chapter 8 An Introduction to Metabolism

Introduction to Metabolism
Metabolism Free Energy

The complexity of metabolism

(greatly simplified)
Only a few hundred of the thousands of metabolic pathways are shown. Insert shows the first two steps in the pathway for the breakdown of glucose
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Metabolism
Sum of all of the biochemical transformations in cell Two divisions Catabolism Anabolism

Metabolism
Catabolism
Degradative reactions Complex molecules are broken down to simple molecules Release of energy to do work in the cell

Metabolism
Anabolism
Synthetic reactions Simple molecules combined to yield complex molecules amino acids combined to form a protein Requires energy

Metabolism
Catabolic and Anabolic pathways often intersect
energy production reactions of catabolism used to drive the energy requiring reactions of anabolism termed Energy Coupling

Energy
Fundamental to all metabolic processes, Important for understanding how the living cell works
The principles that govern energy resources in chemistry, physics, and engineering also apply to bioenergetics, the study of how organisms manage their energy resources.

Metabolism
Energy important in both catabolism and anabolism What is energy?

Metabolism
What is energy? Energy is the capacity to do work.

Energy
Different ways to categorize energy Kinetic Potential

Kinetic Energy
Energy of motion
Moving objects perform work by imparting motion to other matter water moving through dam turns turbines and generates electricity contraction of leg muscles pushes a bicycle pedal moving the bike and rider heat and photons of light - other examples

Potential Energy
Stored energy
Capacity to do work Location or structure of matter water behind a dam (height above the turbine)

Potential Energy
Chemical Energy
a form of potential energy energy stored in molecules as a result of the arrangement of the atoms in the molecules Hydrocarbons in car engine are burned and push pistons - results in movement of car Sugars burned in organisms to supply energy (ATP) to allow muscle movement

Energy Conversion
Energy can be converted from one form to another How is this accomplished?

Energy Conversion
high potential energy
diver on top of platform (1)

potential to kinetic
diver leaps off platform (2)

1 2

potential energy in food the diver ate provided the energy to allow him to climb

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Thermodynamics
Study of energy transformation in matter Three laws of thermodynamics

1st Law of Thermodynamics

Law of Conservation of Energy
The energy of the universe is constant Energy cannot be created or destroyed Can only be transformed from one type to another Plants transform light to chemical energy Conversion is generally reversible

2nd Law of Thermodynamics

All energy transfers or transformations make the universe more disordered (increases entropy).
Entropy = disorder in universe Symbol for entropy is S Change in entropy is S

2nd Law of Thermodynamics

Order may increase locally, but overall in the universe, disorder increases
Increased entropy of the universe takes the form of increasing heat (energy of random molecular motion)

Thermodynamics
In most energy transformations, ordered forms of energy are converted at least partly to heat.
Automobiles convert only 25% of the energy in gasoline into motion; the rest is lost as heat.

Thermodynamics
Living cells unavoidably convert organized forms of energy to heat.
The metabolic breakdown of food ultimately is released as heat even if some of it is diverted temporarily to perform work for the organism.

Heat is energy in its most random state.

Thermodynamics
Combining the two laws of thermodynamics, the quantity of energy is constant, but the quality of energy is not.

Thermodynamics of Living Organisms

Living organisms do not violate the second law of thermodynamics.
Organisms are open systems and take in organized energy like light or organic molecules and replace them with less ordered forms, especially heat.

Thermodynamics of Living Organisms

Organisms show an increase in complexity as they develop
Consistent with the 2nd law as long as the total entropy of the universe increases. Organisms are islands of low entropy (low disorder) in an increasingly random universe.

Free Energy
Reactions can be spontaneous or nonspontaneous
Spontaneous can occur without outside help increase the stability of a system Nonspontaneous: can only occur if energy is added decrease the stability of a system

Free Energy
Can measure the spontaneity of a system using the concept of Free Energy
also called Gibbs Free Energy Josiah Willard Gibbs, American scientist see: http://www.uh.edu/engines/epi119.htm free because it is available to do work

Gibbs Free Energy, G

the portions of a systems energy that is able to perform work when temperature is uniform throughout the system A measure of the instability of the system

Gibbs Free Energy

Symbolized by the letter G free energy (G) in a system is related to the total energy (H) and its entropy (S)

G = H - TS
where T is temperature in Kelvin units

Gibbs Free Energy

A measure of the instability of a system.
Systems that are high in free energy are unstable, e.g. compressed springs, separated charges tend to move toward a more stable state one with less free energy.

Gibbs Free Energy

Systems that tend to change spontaneously are those that have high energy, low entropy, or both. In any spontaneous process, the free energy of a system decreases.

Gibbs Free Energy, G

Change in free energy of a system = G
= Greek letter delta signifies difference or change in

G = Gfinal - Gstarting point G = H - TS

G is negative for a spontaneous reaction
starting energy is > than final energy, so some is available for performing work

Gibbs Free Energy, G

Most chemical reactions are reversible G = 0 at equilibrium
rates of forward and reverse reactions are equal - no change in concentration of reactants and products no work can be done

Fig 8.5 Free energy and work

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Gibbs Free Energy and Metabolism

Chemical Reactions
Exergonic reaction net release of free energy G is negative Endergonic reaction requires energy from its environment G is positive

Fig 8.6a Gibbs Free Energy Change in Exergonic Reactions

net release of free energy

G is negative

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Gibbs Free Energy and Metabolism

Exergonic reaction
magnitude of G is the maximum amount of work the reaction can perform

C6H12O6 + 6O2

6CO2 + 6H2O

Net reaction for Cellular Respiration G = -686 kcal/mole (-2870 kJ/mol)

Fig 8.6b Gibbs Free Energy Change in Endergonic Reactions

net input of free energy required

G is positive non-spontaneous

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Gibbs Free Energy and Metabolism

Cellular respiration releases 686 kcal Photosynthesis, the reverse reaction, must require an equivalent investment of energy.
Delta G = + 686 kcal / mol. Photosynthesis is steeply endergonic, powered by the absorption of light energy

Gibbs Free Energy and Metabolism

Reactions in closed systems eventually reach equilibrium and can do no work.
A cell that has reached metabolic equilibrium has a G = 0 and is dead!

Metabolic dis-equilibrium is one of the defining features of life.

Fig 8.7a Hydraulic analogy for a free energy gradient

Equilibrium (DEATH)

a) electric energy is generated only by falling water

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b) no energy generated at equilibrium

Fig 8.7b Hydraulic analogy for a free energy gradient

Cells maintain disequilibrium because they are open with a constant flow of material in and out of the cell. A cell continues to do work throughout its life
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Fig 8.7c Hydraulic analogy for a free energy gradient

A catabolic reaction in a cell releases free energy in a series of reactions.

Some reversible reactions of respiration are constantly pulled in one direction as the product of one reaction does not accumulate, but becomes the reactant in the next step
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Fig 8.8 ATP Structure

Source of energy for cellular work

ATP

* *
* = high energy phosphates
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ATP Hydrolysis
Hydrolysis of the end phosphate group forms adenosine diphosphate [ATP -> ADP + Pi] and releases 7.3 kcal/mole of ATP under standard conditions. G in the cell is about -13kcal/mole
Fig 8.9

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ATP - the energy molecule

Phosphate bonds
referred to as high-energy phosphate bonds these are actually fairly weak covalent bonds unstable however and their hydrolysis yields energy as the products are more stable. Each of the three phosphate groups has a negative charge Their repulsion contributes to the instability of this region of the ATP molecule.

ATP - the energy molecule

Energy from the hydrolysis of ATP is coupled directly to endergonic processes by transferring the phosphate group to another molecule.
This molecule is now phosphorylated and is now more reactive.

Fig 8.10 Energy coupling by phosphate transfer

Overall reaction has a negative G, so it is spontaneous.

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Regeneration of ATP
After hydrolysis of ATP, cell needs to regenerate it by adding PO4 to ADP
Energy for this comes from catabolic reactions in the cell Endergonic reaction G = 7.3kcal/mole

Fig 8.12 The ATP Cycle Regeneration of ATP

G = -7.3 kcal/mol

G = +7.3 kcal/mol

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Cellular Work
Three main kinds of work
1) Mechanical work beating of cilia & flagella, muscle contraction 2) Transport work pumping of substances across membranes 3) Chemical work getting non-spontaneous reactions to go

Fig 8.11 Mechanical Work

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Fig 8.11b Transport Work

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Fig 8.11c Chemical Work

Copyright 2005 Pearson Education, Inc., publishing as Benjamin Cummings