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BCM301


Biochemistry II


    Chapter 1:
     Bioenergetics


By: Zatilfarihiah Rasdi
By the end of the lecture, students
    should be able to know/define/state:

   The first and second law of thermodynamics
   The enthalpy, entropy and free energy
   Exothermic, endothermic, exergonic and endergonic
    reactions
   Coupled reactions

     Do revise this topic and refer to
      textbook of biochemistry!!!
A Review: Thermodynamic
            Principles
   Living things require a continuous throughput of
    energy. eg. Photosynthesis process – plants convert
    radiant energy from the Sun, the primary energy
    source for life on Earth, to the chemical energy of
    carbohydrates and other organic substances.
   The plants/ animals that eat them, then metabolize
    these substances to power such functions as the
    synthesis of biomolecules, the maintenance of
    concentration gradients and the movement of
    muscles.
   These processes transform the energy to heat, which is
    dissipated to the environment and must be devoted to the
    acquisition and utilization of energy.
   Thermodynamics (Greek: therme, heat + dynamics,
    power) is a description of the relationships among the
    various forms of energy and how energy affects matter on
    the macroscopic as opposed to the molecular level.
   With a knowledge of thermodynamics we can determine
    whether a physical process is possible. Thermodynamics
    is essential for:
       understanding why macromolecules fold to their native
        conformations
       how metabolic pathways are designed, why molecules cross
        biological membranes
       how muscles generate mechanical force
1. First Law of Thermodynamics:
               Energy Is Conserved
   A system is defined as that part of the universe that is of
    interest, such as reaction vessel or an organism; the rest
    of the universe is known as the surroundings.
   A system is said to be open, closed or isolated according
    to whether or not it can exchange matter and energy
    with its surroundings, only energy.
   Living organisms, which take up nutrients, release waste
    products and generate work and heat (open system).
   If an organism were sealed inside an uninsulated box, it
    would, together with the box, constitute a closed
    system.
   If the box perfectly insulated, the system would be
    isolated.
   Processes in which a negative q, are known as exothermic processes (Greek: exo, out of); those in which the

   The SI unit of energy, the joule (J), is steadily replacing the calorie (cal) in modern scientific usage.
Voet Biochemistry 3e Page 52
© 2004 John Wiley & Sons, Inc.




                                              Constants.
                                 Table 3-1 Thermodynamic Units and
a. State functions are independent of the path a systems
       follow
    • Experiments have invariably demonstrated that the energy of a
      system depends only on its current properties or state, not on how
      it reached that state.


B. Enthalpy
-     any combination of only state functions must also be a state
      function. One such combination, known as enthalpy (Greek: to
      warm in) is defined
                           H = U + PV
      where V is the volume and P is its pressure.
     is a particularly convenient quantity with which to describe
      biological systems because under constant pressure, a condition
      typical of most biochemical processes, the enthalpy change
      between the initial and final states of a process, ∆H, is the easily
      measured heat that it generates or absorbs.
   In general, the change of enthalpy in any
    hypothetical reaction pathway can be
    determined from the enthalpy change in any
    other reaction pathway between the same
    reactants and products.
1. Second Law of
     Thermodynamics: The universe
    tends toward maximum disorder

   Spontaneous processes are characterized by the
    conversion of order ( in this case the coherent motion
    of the swimmer’s body) to chaos ( here the random
    thermal motion of the water molecules)
   The 2nd law of thermodynamics expresses this
    phenomenon, provide the criterion for determining
    whether a process is spontaneous.
A.   Spontaneity and disorder
    The spontaneous processes occur in directions
     that increase the overall disorder of the universe
     that is, of the systems and its surroundings.
    Disorder, in this context, is defined as the
     number of equivalent ways, W, of arranging the
     components of the universe.
    (Note: Find the equation that involved with W)
Page 53




 Figure 3-1 Two bulbs of equal volumes connected
                 by a stopcock.
B. Entropy
 In a chemical systems, W, the number of equivalent ways of
  arranging a system in a particular state, is usually
  inconveniently immense.
 In order to be able to deal with W more easily, we define, as

  Ludwig Boltzman in 1877, a quantity known as entropy
  (Greek: en, in + trope, turning):
                       S = kB ln W
  that increases with W but in more manageable way. Here kB is
  the Boltzman constant. Eg. For twin bulb system, S = kBN ln 2,
  so the entropy of the system in its most probable state is
  proportional to the number of gas molecules contains.
Note: Entropy is a state function because it depends only on the
  parameters that describe a state.
   The conclusions based on the twin-bulb apparatus
    may be applied to explain, why blood transports
    between the lungs and the tissues. Solutes in
    solution behave analogously to gases in that they
    intend to maintain a uniform concentration
    throughout their occupied volume – this is their
    most probable arrangement.
   In the lungs-concentration of O2 is higher than in
    venous blood passing through them, more O2 enters
    the blood than leaves it. On the other hand, in the
    tissues- where the O2 concentration is lower than in
    arterial blood, there is net diffusion of O2 from
    blood to the tissues.
Figure 3-3    Relationship of entropy and temperature.
  The structure of water or any other substance becomes
increasingly disordered, that is, its entropy increases, as its
                   temperature rises.
3. Free energy change, ∆G – indicator of spontaneity
• Thermodynamic view: metabolism is an energy
   transforming process whereby catabolism provides
   energy for anabolism.
• What is energy?- “the capacity to cause or
   undergo change”
• Cell and organisms are able to harness forms of
   energy and convert them to other suitable forms to
   support movement, active transport and
   biosynthesis.
   The most important medium of energy exchange
    is ATP – “universal carrier of biological energy”
   Fundamental concept of metabolism:
    i. exergonic – the overall process of catabolism
    releases energy (spontaneous)
    ii. endergonic – the overall process of anabolism
    requires nergy input (nonspontaneous)
   Goal of thermodynamic: to predict the spontaneity
    of a process or reaction. The most useful
    thermodynamic terms is free energy, G or known
    as Gibbs free energy.
   G is an indicator of the energy available from the
    reaction to do work;composed of two components,
    enthalpy (H) and entropy (S).
G = H – TS…………………………….(1)

     where T = temperature in Kelvin (K)
     units for G = joules/mol or kJ/mol
∆G = ∆H –T ∆S……………………....(2)

whether a reaction is spontaneous may be predicted from the following
values of ∆G:

If        ∆G < 0 energy is released;reaction is spontaneous and exergonic
          ∆G = 0 reaction is at equilibrium
          ∆G > 0 energy is required;reaction is nonspontaneous and
                 endergonic

Note: it is very difficult to measure ∆G for a biochemical reaction because
the cellular concentrations of the reactants are very small and hard to
determine experimentally. In order to calculate the energy associated with
biochemical reactions, we must resort to the measurement under a set of
standard.
Standard Free Energy Change, ∆G°’
 This section focus on the most important energy

  molecule, ATP.
 The breakdown of ATP must be exergonic reaction, but

  what is the quantitative amount of energy released under
  std. conditions?
             ATP      ADP + Pi + energy

In your introductory chemistry courses, std. conditions for solute
   reactions were defined as:
   1 atm of pressure, 25°C and initial and products concentration of 1
   M. (but in biochemical process) + condition of a pH of 7        the
   modified ∆G°’.
Table 3-2 Variation of Reaction Spontaneity
                                  (Sign of ∆ G) with the signs of ∆ H and ∆ S.
© 2004 John Wiley & Sons, Inc.
Voet Biochemistry 3e Page 56
4. Chemical equilibria
   The entropy (disorder) of a substance increases with its volume.
    eg. Twin-bulb apparatus – a collection of gas molecules occupied
    all of the volume available to it, maximizes its entropy. Entropy is
    a function of concentration.
   If entropy varies with concentration, so do free energy. The free
    energy change of chemical reaction depends on the concentrations
    of both its reactants and products. eg enzymatic reactions which
    needs substrates (reactants) and on the metabolic demand for their
    products.
   The equilibrium constant of a reaction may therefore be
    calculated from standard free energy data and vice versa.

Note: For more information on equilibrium constants, students may
  refer to textbook and reference book of Biochemistry.
Voet Biochemistry 3e Page 57
© 2004 John Wiley & Sons, Inc.
                                 Table 3-3 Variation of Keq with ∆ G° at 25°C.
Table 3-4 (top) Free Energies of Formation of
           Some Compounds of Biochemical Interest.
Page 58
Table 3-4 (middle) Free Energies of
                                 Formation of Some Compounds of
                                        Biochemical Interest.
© 2004 John Wiley & Sons, Inc.
Voet Biochemistry 3e Page 58
Table 3-4 (bottom) Free Energies of
                                 Formation of Some Compounds of
                                        Biochemical Interest.
© 2004 John Wiley & Sons, Inc.
Voet Biochemistry 3e Page 58
A.   Coupled reactions
    The additivity of free energy changes allows an
     endergonic reaction to be driven by an exergonic
     reaction under the proper conditions.
     (thermodynamic basis for the operation of the
     metabolic pathways since most of these reaction
     sequences comprise endergonic as well as
     exergonic reactions.
     (1)     A+B          C+D          ∆G1
     (2)     D+E           F+G         ∆G2
   If ∆G1 ≥ 0, reaction (1) will not occur spontaneously.
   However, if ∆G2 is sufficiently exergonic so that ∆G1 + ∆G2 < 0,
    then although the equilibrium concentration of D in reaction (1)
    will be relatively small, it will be larger than that in reaction (2).
    As reaction (2) converts D to product, reaction (1) will operate in
    the forward direction to replenish the equilibrium concentration of
    D.
   The highly exergonic reaction (2) therefore drives the endergonic
    reaction (1), and the two reactions are said to be coupled through
    their common intermediate D.
   These coupled reactions proceed spontaneously can also be seen
    by summing reactions (1) and (2) to yield overall reaction
    (3)           A+B+E           C+F+G                    ∆G3

    As long as the overall pathway (reaction sequence) is exergonic, it
    will operate in the forward direction. Thus, the free energy of
    ATP hydrolysis, a highly exergonic process, is harnessed to drive

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Ch06 b
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Ch06
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Ch05
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Ch04 b
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Ch04
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Ch03 cont.
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Ch01 cont.
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Ch01 bcm 311

  • 1. BCM301 Biochemistry II Chapter 1: Bioenergetics By: Zatilfarihiah Rasdi
  • 2. By the end of the lecture, students should be able to know/define/state:  The first and second law of thermodynamics  The enthalpy, entropy and free energy  Exothermic, endothermic, exergonic and endergonic reactions  Coupled reactions Do revise this topic and refer to textbook of biochemistry!!!
  • 3. A Review: Thermodynamic Principles  Living things require a continuous throughput of energy. eg. Photosynthesis process – plants convert radiant energy from the Sun, the primary energy source for life on Earth, to the chemical energy of carbohydrates and other organic substances.  The plants/ animals that eat them, then metabolize these substances to power such functions as the synthesis of biomolecules, the maintenance of concentration gradients and the movement of muscles.
  • 4. These processes transform the energy to heat, which is dissipated to the environment and must be devoted to the acquisition and utilization of energy.  Thermodynamics (Greek: therme, heat + dynamics, power) is a description of the relationships among the various forms of energy and how energy affects matter on the macroscopic as opposed to the molecular level.  With a knowledge of thermodynamics we can determine whether a physical process is possible. Thermodynamics is essential for:  understanding why macromolecules fold to their native conformations  how metabolic pathways are designed, why molecules cross biological membranes  how muscles generate mechanical force
  • 5. 1. First Law of Thermodynamics: Energy Is Conserved  A system is defined as that part of the universe that is of interest, such as reaction vessel or an organism; the rest of the universe is known as the surroundings.  A system is said to be open, closed or isolated according to whether or not it can exchange matter and energy with its surroundings, only energy.  Living organisms, which take up nutrients, release waste products and generate work and heat (open system).  If an organism were sealed inside an uninsulated box, it would, together with the box, constitute a closed system.  If the box perfectly insulated, the system would be isolated.
  • 6. Processes in which a negative q, are known as exothermic processes (Greek: exo, out of); those in which the  The SI unit of energy, the joule (J), is steadily replacing the calorie (cal) in modern scientific usage.
  • 7. Voet Biochemistry 3e Page 52 © 2004 John Wiley & Sons, Inc. Constants. Table 3-1 Thermodynamic Units and
  • 8. a. State functions are independent of the path a systems follow • Experiments have invariably demonstrated that the energy of a system depends only on its current properties or state, not on how it reached that state. B. Enthalpy - any combination of only state functions must also be a state function. One such combination, known as enthalpy (Greek: to warm in) is defined H = U + PV where V is the volume and P is its pressure.  is a particularly convenient quantity with which to describe biological systems because under constant pressure, a condition typical of most biochemical processes, the enthalpy change between the initial and final states of a process, ∆H, is the easily measured heat that it generates or absorbs.
  • 9. In general, the change of enthalpy in any hypothetical reaction pathway can be determined from the enthalpy change in any other reaction pathway between the same reactants and products.
  • 10. 1. Second Law of Thermodynamics: The universe tends toward maximum disorder  Spontaneous processes are characterized by the conversion of order ( in this case the coherent motion of the swimmer’s body) to chaos ( here the random thermal motion of the water molecules)  The 2nd law of thermodynamics expresses this phenomenon, provide the criterion for determining whether a process is spontaneous.
  • 11. A. Spontaneity and disorder  The spontaneous processes occur in directions that increase the overall disorder of the universe that is, of the systems and its surroundings.  Disorder, in this context, is defined as the number of equivalent ways, W, of arranging the components of the universe.  (Note: Find the equation that involved with W)
  • 12. Page 53 Figure 3-1 Two bulbs of equal volumes connected by a stopcock.
  • 13. B. Entropy  In a chemical systems, W, the number of equivalent ways of arranging a system in a particular state, is usually inconveniently immense.  In order to be able to deal with W more easily, we define, as Ludwig Boltzman in 1877, a quantity known as entropy (Greek: en, in + trope, turning): S = kB ln W that increases with W but in more manageable way. Here kB is the Boltzman constant. Eg. For twin bulb system, S = kBN ln 2, so the entropy of the system in its most probable state is proportional to the number of gas molecules contains. Note: Entropy is a state function because it depends only on the parameters that describe a state.
  • 14. The conclusions based on the twin-bulb apparatus may be applied to explain, why blood transports between the lungs and the tissues. Solutes in solution behave analogously to gases in that they intend to maintain a uniform concentration throughout their occupied volume – this is their most probable arrangement.  In the lungs-concentration of O2 is higher than in venous blood passing through them, more O2 enters the blood than leaves it. On the other hand, in the tissues- where the O2 concentration is lower than in arterial blood, there is net diffusion of O2 from blood to the tissues.
  • 15. Figure 3-3 Relationship of entropy and temperature. The structure of water or any other substance becomes increasingly disordered, that is, its entropy increases, as its temperature rises.
  • 16. 3. Free energy change, ∆G – indicator of spontaneity • Thermodynamic view: metabolism is an energy transforming process whereby catabolism provides energy for anabolism. • What is energy?- “the capacity to cause or undergo change” • Cell and organisms are able to harness forms of energy and convert them to other suitable forms to support movement, active transport and biosynthesis.
  • 17. The most important medium of energy exchange is ATP – “universal carrier of biological energy”  Fundamental concept of metabolism: i. exergonic – the overall process of catabolism releases energy (spontaneous) ii. endergonic – the overall process of anabolism requires nergy input (nonspontaneous)  Goal of thermodynamic: to predict the spontaneity of a process or reaction. The most useful thermodynamic terms is free energy, G or known as Gibbs free energy.  G is an indicator of the energy available from the reaction to do work;composed of two components, enthalpy (H) and entropy (S).
  • 18. G = H – TS…………………………….(1) where T = temperature in Kelvin (K) units for G = joules/mol or kJ/mol ∆G = ∆H –T ∆S……………………....(2) whether a reaction is spontaneous may be predicted from the following values of ∆G: If ∆G < 0 energy is released;reaction is spontaneous and exergonic ∆G = 0 reaction is at equilibrium ∆G > 0 energy is required;reaction is nonspontaneous and endergonic Note: it is very difficult to measure ∆G for a biochemical reaction because the cellular concentrations of the reactants are very small and hard to determine experimentally. In order to calculate the energy associated with biochemical reactions, we must resort to the measurement under a set of standard.
  • 19. Standard Free Energy Change, ∆G°’  This section focus on the most important energy molecule, ATP.  The breakdown of ATP must be exergonic reaction, but what is the quantitative amount of energy released under std. conditions? ATP ADP + Pi + energy In your introductory chemistry courses, std. conditions for solute reactions were defined as: 1 atm of pressure, 25°C and initial and products concentration of 1 M. (but in biochemical process) + condition of a pH of 7 the modified ∆G°’.
  • 20. Table 3-2 Variation of Reaction Spontaneity (Sign of ∆ G) with the signs of ∆ H and ∆ S. © 2004 John Wiley & Sons, Inc. Voet Biochemistry 3e Page 56
  • 21. 4. Chemical equilibria  The entropy (disorder) of a substance increases with its volume. eg. Twin-bulb apparatus – a collection of gas molecules occupied all of the volume available to it, maximizes its entropy. Entropy is a function of concentration.  If entropy varies with concentration, so do free energy. The free energy change of chemical reaction depends on the concentrations of both its reactants and products. eg enzymatic reactions which needs substrates (reactants) and on the metabolic demand for their products.  The equilibrium constant of a reaction may therefore be calculated from standard free energy data and vice versa. Note: For more information on equilibrium constants, students may refer to textbook and reference book of Biochemistry.
  • 22. Voet Biochemistry 3e Page 57 © 2004 John Wiley & Sons, Inc. Table 3-3 Variation of Keq with ∆ G° at 25°C.
  • 23. Table 3-4 (top) Free Energies of Formation of Some Compounds of Biochemical Interest. Page 58
  • 24. Table 3-4 (middle) Free Energies of Formation of Some Compounds of Biochemical Interest. © 2004 John Wiley & Sons, Inc. Voet Biochemistry 3e Page 58
  • 25. Table 3-4 (bottom) Free Energies of Formation of Some Compounds of Biochemical Interest. © 2004 John Wiley & Sons, Inc. Voet Biochemistry 3e Page 58
  • 26. A. Coupled reactions  The additivity of free energy changes allows an endergonic reaction to be driven by an exergonic reaction under the proper conditions. (thermodynamic basis for the operation of the metabolic pathways since most of these reaction sequences comprise endergonic as well as exergonic reactions. (1) A+B C+D ∆G1 (2) D+E F+G ∆G2
  • 27. If ∆G1 ≥ 0, reaction (1) will not occur spontaneously.  However, if ∆G2 is sufficiently exergonic so that ∆G1 + ∆G2 < 0, then although the equilibrium concentration of D in reaction (1) will be relatively small, it will be larger than that in reaction (2). As reaction (2) converts D to product, reaction (1) will operate in the forward direction to replenish the equilibrium concentration of D.  The highly exergonic reaction (2) therefore drives the endergonic reaction (1), and the two reactions are said to be coupled through their common intermediate D.  These coupled reactions proceed spontaneously can also be seen by summing reactions (1) and (2) to yield overall reaction (3) A+B+E C+F+G ∆G3 As long as the overall pathway (reaction sequence) is exergonic, it will operate in the forward direction. Thus, the free energy of ATP hydrolysis, a highly exergonic process, is harnessed to drive