Chapter 9 CELLULAR RESPIRATION:

HARVESTING CHEMICAL ENERGY

Summary of Chapter 9, BIOLOGY, 9
TH
ED Campbell, by J.B. Reee et al. !"##.
Living is work.
Metabolic reactions require energy transformations.
Metabolism allows the organism to move, grow, heal, reproduce, etc.
Cells make polymers, pump substances across membranes at the expense of energy, reproduce,
etc.
xception of thermal vents at the bottom of oceans, the source of all energy used in the
biosphere is the sun.
PRINCIPLES OF ENERGY HARVEST
Catabolism is the splitting of large molecules into smaller ones. nergy is released in the
process.
!erobic respiration, anaerobic respiration, fermentation.
"rganic compounds store energy in their chemical bonds.
#ith the help of en$ymes, the cell breaks down these organic compounds in a systematic fashion
and makes the chemical energy %potential energy& stored in those bonds, available to anabolic
reactions.
!t any step of the degradation of organic molecules into simpler one, there is a loss of some
energy in the form of heat.
'econd law of thermodynamics( transfer of energy is never )**+ efficient.
Aerobic resiratio! is the most efficient method of degrading energy rich organic compounds.
"xygen is used during cellular respiration to oxidi$e the organic compounds.
Ferme!tatio! is the partial degradation of organic compounds.
,ermentation is conducted in the absence of oxygen.
-he usual fuel molecule consumed in cellular respiration is glucose, but proteins, fats and other
carbohydrates are also used.
-he breakdown of glucose is e"er#o!ic, releasing $%&% 'cal(mole of glucose or ./,01* k23mole
of glucose.
-he products of these reactions have less energy than the reactants.
A)ENOSINE TRIPHOSPHATE* ATP
4n living cell, energy is temporarily stored in !-5.
!-5 donates energy through the transfer of a phosphate group.
!-5 is formed by the +os+or,latio! of !65. -his is an endergonic reaction that requires
energy input.
5hosphorylation occurs when a phosphate group is transferred to some other compound.
!-5 is the link between exergonic catabolic reactions and endergonic anabolic reactions.
RE)O- REACTIONS: O-I)ATION AN) RE)UCTION
nergy can be transferred through the transfer of a phosphate group.
-he loss of electrons by one substance is called o"i.atio!.
-he acceptance of electrons by a molecule is called re./ctio!.
Atte!tio!0: !dding electrons is called reduction because this re$ue% the &umber of po%'t'(e
har)e% in the accepting molecule or atom.
nergy can also be transferred through the transfer of electrons.
lectrons released through oxidation cannot exist in the free state in the cell.
very oxidation reaction must be accompanied by a reduction reaction in which electrons are
accepted by other molecule, ion or atom.
"xidation.reduction reactions are called re.o" reactio!s.
-he electron donor is called the re./ci!# a#e!t.
-he electron acceptor is called the o"i.i1i!# a#e!t.
Changing the degree of electron sharing, rather than losing or gaining electrons, causes the loss
of energy. -his change in electron sharing is also considered a redox reaction.
-he shifting of electrons closer to the very electronegative oxygen %oxidation& causes the loss of
energy that can be put to work.
-he ma7ority of biochemical reduction reactions involve the transfer of a hydrogen atom.
-he hydrogen atom brings with it one electron, therefore, reduction occurs.
"xidation can be defined as the loss of a hydrogen atom, a proton plus an electron.
!erobic respiration is a redox process that transfers hydrogen from sugars to oxygen.
Vale!ce electro!s o2 carbo! a!. +,.ro#e! lose ote!tial e!er#, as t+e, are asse. to t+e
more electro!e#ati3e o",#e!4
-he covalent bonds holding the hydrogen and carbon atoms together lose energy when the
carbon and hydrogen atoms bind to oxygen and the oxygen pulls the electrons closer.
lectrons %negative& closer to the nucleus of oxygen %positive& are more stable8 they have less
energy, and ...
-he released energy is used by the cell to make !-5.
Carbohydrates and fats are excellent energy stores because they have many C . 9 bonds.
Most organisms cataboli$e #l/cose into water and carbon dioxide in order to obtain energy.
C%H56O% 7 %O6 %CO6 7 % H6O 7 e!er#, 8as ATP9
:lucose is oxidi$ed to carbon dioxide( it gives up )/ hydrogen atoms resulting in C"/.
"xygen is reduced to water( it receives )/ hydrogen atoms to make 9/".
-here is a net gain of ; water molecules.
Cellular respiration does not oxidi$e glucose in one step, which would be explosive if done, and
the energy could not be efficiently harnessed in a form available to perform cellular work.
n$ymes lower the activation energy and glucose is slowly oxidi$ed in a series of reactions.
very oxidation reaction must be accompanied by a reduction reaction in which electrons are
accepted by other molecule, ion or atom.
<edox reactions occur simultaneously.
"ften occur in a series of reactions in which electrons are transferred from one
molecule to another.
!t key steps, electrons are stripped from the glucose.
lectron transfers are equivalent to energy transfers.
Most redox reactions involve the transfer of 9 atom, an electron plus a proton.
#hen an electron singly or as part of 9 atom is transferred, it carries with it some of the energy
stored in the chemical bond.
-he electron progressively loses free energy as it is transferred from molecule to molecule.
Carbohydrates and fats are reservoirs of electrons associated with hydrogen
"nly the barrier of activation energy holds back the flood of electrons to a lower energy state.
#ithout this barrier, glucose will combine instantly with oxygen in the atmosphere.
A co!trolle. e"er#o!ic reactio!4 Ste:ise E!er#, Har3est 3ia NA)
7
a!. t+e Electro!
Tra!sort C+ai!4
-he release of energy stored in glucose all at once will be explosive.
Cellular respiration does not transfer all hydrogen atoms in a single explosive step.
:lucose and other fuel molecules are broken down gradually in a series of steps cataly$ed by an
en$yme.
9ydrogen atoms stripped from glucose are not transferred directly to oxygen but to intermediate
9 carriers, =!69 and ,!69/.
)e+,.ro#e!ase en$ymes cataly$e the removal of hydrogen from glucose and their transfer
=!6
>
.
Nicoti!ami.e a.e!i!e .i!/cleoti.e or NA)
7
is a common electron acceptor, therefore, an
oxidi$ing agent. See *'). 9.+ o& pa)e #,,.
?9/ > =!6
>
? > =!69 > 9
>

oxidi$ing oxidi$ed reduced
agent
-wo protons and two electrons %@ / hydrogen atoms& were removed from ?9/, a substrate,
e.g. a sugar.
=!6
>
is an oxidi$ing agent.
-wo electrons and one proton were transferred to NA)
7
4
-he other proton is released to the surroundings as 9
>
.
-he oxidi$ed form of NA)
7
has a positive charge. 4t is an oxidi$ing agent %9 acceptor&.
-he reduced form is neutral, NA)H.
-he energy stored in =!69 is usually transferred to !-5.
Nicoti!e a.e!i!e .i!/cleoti.e +os+ate or NA)P
7
is another acceptor molecule similar to
=!6
>
and form the reduced =!659. 4t is found in plants.
=!659 provides energy to other reactions including some reactions of photosynthesis.
Fla3i! a.e!i!e .i!/cleoti.e or FA), accepts hydrogen atoms and becomes ,!69/.
-he c,toc+romes are proteins a heme prosthetic group that contain iron. -he iron atom accepts
electrons from 9 atoms and transfers these electrons to other compounds.
NA)H and FA)H6 transfer these electrons to oxygen to make water in a series of steps called the
electro! tra!sort c+ai!.
-he electron transport chain is a downhill energy path for the electrons.
-he total amount of energy released by =!69 in the electron transport chain is .AB kcal3mole %.
/// k23mole&.
,ood has high.energy hydrogen atoms %electrons& forming bonds with carbon and at the end of
cellular respiration, these hydrogen atoms %electrons& have donated their energy to !-5,
becoming low.energy electrons and 9
>
that form water by reacting with oxygen, "/.
'CMM!<D( electrons follow this pathE
,ood %glucose& F =!69 F electron transport chain F oxygen
CELLULAR or AERO;IC RESPIRATION
Cellular respiration is an aerobic process. 4t uses oxygen.
See $'a)ram o& pa)e #,-, *'). 9.,.
54 GLYCOLYSIS
4t takes place in the cytosol of the cell.
! glucose molecule %;.C& is converted to two B.C pyruvate molecules.
/!-5 and /=!69 are formed.
S/bstrate le3el +os+or,latio! makes !-5. !n organic molecule donates the phosphate.
=!69 carries the 9 atoms to the electron transport chain to make more !-5.
5yruvate exists in the cell as an anion.
="-4C( no oxygen is needed in glycolysis8 no C"/ if produced.
'CMM!<D( / !-58 this occurs twice per glucose molecule.
'ee page );9. 4M5"<-!=-.
64 FORMATION OF ACETYL COEN<YME A 8CoA49
ach pyruvate enters the matrix of the mitochondrion in eukaryotic cells. 4n prokaryotic cells it
occurs in the cytosol.
"ne molecule of C"/ is produced from the carboxyl group of the pyruvate molecule.
-he remaining two.carbon fragment is oxidi$ed to a /.C acetate, which combines with
coen$yme ! forming acet,l$CoA.
!n en$yme transfer the extracted electrons to =!6
>
. =!69 is produce.
! multien$yme complex cataly$es this step.
Co! is a sulfur.containing compound derived from vitamin G that forms an unstable bond with
the acetyl group and makes it very reactive.
'CMM!<D( )C"/8 )=!69. -his occurs twice per molecule of glucose.
'ee fig. 9.)* on page )1*. 4M5"<-!=-.
=4 >RE;S or CITRIC ACI) CYCLE
!lso known as the Hrebs cycle and tricarboxylic acid %-C!&.
-wo acetyl.Co! produced from one glucose molecule enter the citric acid cycle.
4t takes place in the mitochondrial matrix and a specific en$yme cataly$es each step.
-he acetate group of acetyl.Co! combines with a I.C molecule of oxaloacetate, and forms a
;.C molecule of citrate.
Citrate is recycled to oxaloacetate in a series of reactions.
T:o CO6 molecules are produced in the process
9 atoms are transferred to t+ree NA)H and o!e FA)H6. -otal of ; =!69 and / ,!69/
molecules per glucose.
O!e ATP molecule is formed from each acetate by substrate.level phosphorylation. -otal of /
!-5 per glucose molecule.
'CMM!<D( / C"/8 B =!698 ) ,!69/8 ) !-58 this occurs twice per molecule of glucose.
'ee diagrams fig. 9.)) and fig. 9.)/ on pages )1* and )1). 4M5"<-!=-.
?4 ELECTRON TRANSPORT CHAIN AN) CHEMIOSMOSIS
-he electron transport chain is a series of molecules embedded in the inner membrane of the
mitochondrion. -here are thousands of copies of the -C in the inner membrane.
-hese molecules are proteins with rost+etic #ro/s %non.protein& bound to them.
lectron carriers alternate between oxidi$ed and reduced states. ach carrier passes
electrons to its JdownhillK more electronegative neighbor.
=!69 and ,!69/ carry the electrons transferred in the previous stages to a chain of electron
acceptors.
!s electrons are transferred from one acceptor to another, some of their energy is used to
pump 9
>
, protons, across the inner mitochondrial membrane, forming a proton gradient.
9igh proton concentration in the intermembrane space and low concentration in the matrix.
6uring chemiosmosis, the proton gradient is used to synthesi$e !-5.
-he synthesis of !-5 from !65 and 5 during the -C is called o"i.ati3e +os+or,latio!.
"xygen is the final electron acceptor.
! maximum of about BI . B0 !-5s molecules are synthesi$ed from one glucose molecule.
'ee fig. 9.)B, on page )1B. 4M5"<-!=-.
5rosthetic groups are cofactors that tightly bind to proteins in order to activate the en$yme.
CHEMIOSMOSIS
T+e #ra.ie!t across a membra!e /se. to .ri3e t+e s,!t+esis o2 ATP is calle.
c+emiosmosis4
:lycolysis and the Hrebs cycle account for only four !-5 molecules. !t this point, most of the
energy extracted from glucose is found in =!69 and ,!69/.
Str/ct/ral .etails o2 t+e ETC
-he electron transport chain %-'& is located in the inner mitochondrial membrane.
-he folded inner membrane of the mitochondrion provides the space for thousands of copies of
the chain.
Most components of the chain are proteins. "ne of the molecules involved is a lipid called
/bi@/i!o!e, L.
-ightly bound to these proteins are prosthetic groups, non.protein components essential for the
catalytic function of certain en$ymes.
-he proteins are called c,toc+romes and have a +eme group as a prosthetic group. 9eme
groups contain an iron atom.
-hese prosthetic groups alternate between reduced and oxidi$ed states as they accept and
donate electrons.
Most of these electron carriers %cytochromes& are grouped into multiprotein complexes.
More on electron carriers( http(33rpi.edu3dept3bcbp3molbiochem3MG#eb3mb)3part/3redox.htm
-he inner membrane of the mitochondrion or prokaryotic plasma membrane contains many
copies of a protein complex called ATP s,!t+ase.
!-5 synthase is an ion pump acting in reverse8 it pumps 9
>
from the intermembrane
space into the mitochondrial matrix.
-he -C pumps 9
>
from the mitochondrial matrix into the intermembrane space.
-he en$yme complex ATP s,!t+ase is made of three parts(
). ! cylindrical rotor within the membrane.
/. ! rod connecting the cylindrical rotor to the knob.
B. ! knob protruding into the matrix.
'ee diagrams on pages )1I and )1A, figs. 9.)I and 9.)A. 4M5"<-!=-.
E!er#, co/li!# mec+a!ism
-his model is widely accepted now and it was proposed in )9;) by the Gritish biochemist 5eter
Mitchell.
lectrons fall to successively lower energy levels as they are passed along the four protein
complexes of the -C.
'ome of the energy released is used by three of the complexes to pump protons out of the matrix
and across the inner mitochondrial membrane into the intermembrane space.
! proton gradient is established across the inner mitochondrial membrane with the higher
concentration %lower p9& in the intermembrane space between the inner membrane and the outer
membrane of the mitochondrion, and the lower concentration %higher p9& in the matrix.
5rotons are pumped across the membrane by three electron transport complexes.
-he gradient is a form of potential energy. -he electrons in the intermembrane space have
thermodynamic tendency to flow back to the matrix8 to flow from the area of higher concentration
to the area of lower concentration.
nergy is spent in order to maintain the proton gradient.
6iffusion of the protons back into the matrix occurs through specific channels formed by the
en$yme complex ATP s,!t+ase* a transmembrane protein complex.
Str/ct/re o2 t+e ATP s,!t+ase( 4t is a protein complex made of many polypeptide subunits
arranged in four main parts, each made of multiple polypeptides.
). -he rotor embedded in the inner mitochondrial membrane.
/. -he knob that protrudes into the mitochondrial matrix.
B. !n internal rod extending from the rotor into the knob.
I. -he stator anchored next to the rotor that holds the knob stationary.
IMPORTANT( 'tudy ,ig. 9.)I on page )1I. 9ow !-5 synthase is made and works.
lectrons flowing between the stator and rotor cause the rotor and its attached rod to rotate.
-he rotating rod causes conformational changes in the stationary knob activating three catalytic
sites to promote the combination of !65 and 5i to produce !-5.
6iffusion of protons back into the matrix through the !-5 synthase across the membrane is
exergonic and it provides the energy for !-5 synthesis. 'ome energy is dissipated as heat.
See f'). 9.#+
!lso( http(33www.bio.davidson.edu3Courses3Molbio3Mol'tudents3spring/**B3Gennett3protein).htm
-he 9
>
gradient has the capacity to do work and is called a roto!$moti3e 2orce.
!-5 is produced through the phosphorylation of !65.
Chloroplasts use chemiosmosis to make !-5.
5rokaryotes lack mitochondria. -hey produce 9
>
across the plasma membrane.
-he proton.motive force is used to make !-5, and to pump nutrients into the cell and wastes out
of the cell.
,or a detail explanation of how the complexes of the -C work see the following animation(
http(33www.brookscole.com3chemistryMd3templates3studentMresources3sharedMresources3animatio
ns3oxidative3oxidativephosphorylation.swf
ATP YIEL)
"ne molecule of glucose produces...
:lycolysis( 6 ATP and 6 NA)H
/ pyruvates / acetyl Co! and 6 NA)H
/ acetyl Co! % NA)H > 6FA)H6 > 6ATP
-he total is )* =!69, I !-5 and / ,!69/.
ach =!69 generates about B !-5s producing a total of 6& to =A ATP in the -C.
ach ,!69/ produces / !-5s giving a total of ? ATPs per glucose.
? ATPs were produced directly during glycolysis and citric cycle through substrate level
phosphorylation.
-he grand total is =% to =& ATPs4 -his is only a! estimate.
=!69 generated in the cytosol during glycolysis cannot pass through the membranes into the
mitochondrial matrix. -here is a shuttle system that transports the electrons of the cytosol =!69
to molecules of =!6
>
or ,!6 in the matrix.
-he number of !-5 produced will depend on the recipient of the electrons, =!6
>
or ,!6.
-he transfer of two electrons from =!69 to oxygen usually results in the production of three !-5
molecules in bacteria.
ukaryotes must spend some !-5 in transporting electrons from the cytosol to the matrix and
usually generate less !-5 per glucose molecule than bacteria
-his number, NB0 !-5 molecules, represents about I*+ efficiency in transfer of energy from
glucose to !-5. -he rest is dissipated as heat or used to maintain a stable body temperature.
Gody heat is a byproduct of the exergonic -C.
RELATE) META;OLIC PROCESSES
ANAERO;IC RESPIRATION
'ome types of bacteria that live in an oxygen.depleted environment in water.logged soil, pond
sediments and animal intestines perform anaerobic respiration.
lectrons are transferred to =!69 and then to the -C, and chemiosmosis makes !-5.
!n i!or#a!ic s/bsta!ce is t+e 2i!al electro! accetor, e.g. ' to '
.

-he final products are C"/, one or more reduced inorganic substances like 9/', and !-5.
-he sulfate reducing bacteria use the sulfate ion at the end of their electron transport
chain instead of oxygen.
4t is anaerobic because it doesnOt use oxygen but there is an -C.
FERMENTATION
-his a!aerobic at+:a, does not involve the electron transport chain.
"xidation is the loss of electrons to !=D acceptor.
:lycolysis is the first part of the pathway.
:lucose is oxidi$ed to two molecules of pyruvate.
=!6
>
is the oxidi$ing agent. -he product is =!69.
/ !-5 are made by s/bstrate$le3el +os+or,latio!.
:lycolysis produces two !-5s whether there is oxygen present or not.
4n an anaerobic environment, =!69 regenerates =!6
>
by transferring electrons and hydrogen to
pyruvate or a derivative of pyruvate.
<emember that in an aerobic environment, =!69 goes to the -C and =!6
>
is
regenerated.
-wo common types of fermentation are(
4n alco+ol 2erme!tatio!, the produced B.C pyruvate is converted to C"/ and the /.C ethyl
alcohol. See *'). 9.#-.a/.
-he =!69 produced in glycolysis is used in the formation of alcohol and C"/ and =!6
>
is
regenerated.
4n lactate or lactic aci. 2erme!tatio! the final product is the B.C lactate and =!69 is oxidi$ed
to =!6
>
. See *'). 9.#-.b/.
-he hydrogen atoms of =!69 are added to pyruvate to make lactate.
"nly / !-5s are produced per glucose molecule.
E3ol/tio!ar, si#!i2ica!ce o2 #l,col,sis.
5rokaryotes appear in the fossil record about B.A billion years ago.
"xygen began to accumulate in the atmosphere about /.1 billion years ago, when cyanobacteria
appeared.
:lycolysis probably evolved in prokaryotes as a means of producing !-5 in an atmosphere
without oxygen.
:lycolysis is the most widespread metabolic pathway, which suggests a very early appearance
before groups began to branch out.
:lycolysis does not require any membrane enclosed organelle suggesting that it evolved before
the appearance of eukaryotic cells.
OTHER NUTRIENTS
:lycolysis and the Hrebs cycle connect many other metabolic pathways.
Pol,sacc+ari.es(
'tarch, glycogen and sucrose provide glucose and fructose for glycolysis.
Protei!s(
!mino acids produced during protein digestion are .eami!ate. and the amino group is
converted to urea.
'ome !! generate pyruvate or oxaloacetate which is converted to glucose.
"thers are converted into one of the reactants in the citric acid chain, e.g. oxaloacetate,
succinate, fumarate, ketoglutarate or to acetyl.Co!.
Fats(
,ats are broken down into glycerol and fatty acids.
:lycerol is converted to glyceraldehyde.B.phosphate, which is one the intermediate compounds
in glycolysis.
,atty acids are oxidi$ed and split into /.C acetyl groups and converted to acetyl.Co! in a process
called .oxidation.
!cetyl.Co! enters the citric acid cycle.
REGULATION OF AERO;IC RESPIRATION
!erobic respiration is controlled through a negative feedback mechanism
-he en$yme phosphofructokinase binds a phosphate to fructose.;.phospate in the early part of
glycolysis and produces fructose.),;.phosphate.
Large amounts of !-5 inactivate the en$yme by !-5 binding to the allosteric site and rendering it
inactive and slowing down !-5 production.
!M5 and !65 activate the en$yme.
5hosphofructokinase can also be inhibited by citrate, one of the Hrebs cycle molecules.
!s citrate accumulates, glycolysis slows down and the supply of acetyl.Co! to the Hrebs cycle
slows down.
S/mmar,(
). Catabolism and anabolism.
!nabolic pathways use energy
Catabolic pathways release energy by oxidi$ing organic molecules.
/. "xidation reduction
:ain and loss of electrons
:ain and loss of a hydrogen atom
Cneven sharing of electrons
B. nergy released in a stepwise mechanism
I. <ole of =!69, ,!69/ and !-5.
A. 'tages of cellular respiration
:lycolysis
Citric !cid Cycle or Hrebs Cycle
lectron -ransport Chain or -C
Conversion of pyruvate to acetyl Co.!
Hnow the reactant and product for each stage8 how many =6!9, ,!9/ and !-5
molecules are produced.
;. Chemiosmosis
#hat is chemiosmosisP
nergy coupling mechanism of the electron transport chain to !-5 synthesis
"xidative phosphorylation( what is itP
!ccounting of !-5 production by cellular respiration( how many !-5 are
produced3glucose molecule
#hat is the difference between substrate.level phosphorylation and oxidative
phosphorylationP
1. ,ermentation and anaerobic respiration
0. #hat is the difference between themP
9. -ypes of fermentation( lactic acid and alcohol.
)*. Catabolism of carbohydrates, proteins and lipids