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08 Metabolism, cell respiration,

and photosynthesis

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Essential ideas Organisms on our planet are
part of a balanced system in
8.1 Metabolic reactions are regulated in response to the cells needs. which the products of the
metabolic processes of one
group of organisms are shared
8.2 Energy is converted to a usable form in cell respiration. by all organisms. There is a
constant interaction amongst
all species. We must be ever-
mindful of this balance and
8.3 Light energy is converted into chemical energy. must continually work towards
its maintenance.

Energy is a topic of discussion every day in our modern world. We talk about the
energy needed to run our modes of transport. We talk about being so tired after a long
day at school that we need a short nap. The need for food becomes essential at times to
regain the energy levels necessary for us to function. This chapter will look at energy in
living systems. The harnessing of energy by plants will be discussed in detail, as well as
how both plants and animals may then release this harnessed energy in a form usable
by the organism in question. The role of enzymes in these energy processes will be
examined first.


8.1 Metabolism Developments in

scientific research
follow improvements in
computing: developments
Understandings: in bioinformatics, such
Metabolic pathways consist of chains and cycles of enzyme-catalysed reactions. as the interrogation of
Enzymes lower the activation energy of the chemical reactions that they catalyse. databases, have facilitated
Enzyme inhibitors can be competitive or non-competitive. research into metabolic
Metabolic pathways can be controlled by end-product inhibition. pathways.

Applications and skills:

Application: End-product inhibition of the pathway that converts threonine to isoleucine.
Application: Use of databases to identify potential new anti-malarial drugs.
Skill: Calculating and plotting rates of reactions from raw experimental results.

Skill: Distinguishing different types of inhibition from graphs at a specified substrate concentration.

Enzyme inhibition should be studied using one specific example for competitive and non-competitive


Your metabolism is the sum of all the chemical reactions that occur within you as
a living organism. The type of reaction that uses energy to build complex organic
molecules from simpler ones is called anabolism. The type of reaction that breaks
down complex organic molecules with the release of energy is called catabolism.
Table 8.1 summarizes anabolic and catabolic reactions.


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08 Metabolism, cell respiration, and photosynthesis

Endergonic reactions are Table 8.1 Anabolic and catabolic reactions

said to occur when the
products of a chemical Anabolic reactions Catabolic reactions
reaction have more
Build complex molecules Break down complex molecules
energy than the reactants
or substrates of the Are endergonic Are exergonic
reaction. Endergonic
reactions tend to Are biosynthetic Are degradative
occur in biosynthetic
reactions in which more Example: photosynthesis Example: cellular respiration
complex molecules
are produced in the
reaction. In contrast, Metabolic pathways
exergonic reactions
occur when the products Almost all metabolic reactions in organisms are catalysed by enzymes. Many of
of a chemical reaction these reactions occur in specific sequences and are called metabolic or biochemical
have less energy than pathways. A very simple, generalized, metabolic pathway is represented in Figure 8.1.
the reactions reactants
or substrates. Exergonic
reactions tend to occur in Figure 8.1 An
degradative reactions or example of a
those reactions in which metabolic pathway.
Substrate A Substrate B Final product
a complex molecule
is broken down into
simpler materials. Each arrow represents a specific enzyme that causes one substrate to be changed to
another, until the final product of the pathway is formed. Some metabolic pathways
Biological processes and
consist of cycles of reactions instead of chains of reactions. Others involve both cycles
chemical pathways are and chains of reactions. Cell respiration and photosynthesis were discussed in Chapter
usually quite complex. 2, and both are complex pathways with chains and cycles of reactions. Metabolic
When scientists attempt pathways are usually carried out in designated compartments of the cell where the
to explain these complex
reactions, they usually
necessary enzymes are clustered and isolated. The enzymes required to catalyse every
break them down into reaction in these pathways are determined by the cells genetic makeup.
smaller, intermediate
steps. These intermediate
steps are then carefully
researched and imitated.
Induced-fit model of enzyme action
The hope is that eventually Enzymesubstrate specificity was discussed in Section 2.5. Enzyme specificity is
an understanding of made possible by enzyme structure. Enzymes are very complex protein molecules
the complete process
with high molecular weights. The higher levels of protein structure allow enzymes to
is obtained. There are
many subjects where form unique areas, such as the active site. The active site is the region on the enzyme
knowledge is gained in a that binds to a particular substrate or substrates. This binding results in the reaction
similar manner. Discuss occurring much faster than would be expected without the enzyme.
some examples. Is this
manner of understanding In the 1890s Emil Fischer proposed the lock-and-key model of enzyme action. He
complex concepts always suggested that substrate molecules fit like a key into a rigid section of the enzyme
successful? What are some
lock. At the time this model provided a good explanation of the specificity of enzyme
of the limitations of this
approach? action. However, as knowledge about enzyme action has increased, Fischers model
has been modified.

Enzymes are globular

It is now obvious that many enzymes undergo significant changes in their
proteins that, as conformation when substrates combine with their active site. The accepted new model
a minimum, have for enzyme action is called the induced-fit model. A good way to visualize this model
the tertiary level of of enzyme action is to think of a hand and glove, the hand being the substrate and the
glove being the enzyme. The glove looks a bit like the hand. However, when the hand
is actually placed in the glove, there is an interaction that results in a conformational
change of the glove, thus providing an induced fit.

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The conformational changes and induced fit are the result of changes in the R-groups
In 1958, Daniel Koshland
of the amino acids at the active site of the enzyme, as the enzyme interacts with the used a larger body of
substrate or substrates. knowledge than had been
available to Fischer, to
present the induced-fit
Activation energy model.

When talking about enzyme action, we always refer to activation energy (AE).
Activation energy is best understood as the energy necessary to destabilize the existing
chemical bonds in the substrate of an enzymesubstrate catalysed reaction. Enzymes
work by lowering the activation energy required (see Figure 8.2). This means that the
enzymes cause chemical reactions to occur faster because they reduce the amount of
energy needed to bring about the chemical reaction.
It is important to note that, even though enzymes lower the activation energy of a
particular reaction, they do not alter the proportion of reactants to products.
energy supplied

uncatalysed Figure 8.2 Enzymes accelerate

activation energy reactions by lowering the
catalysed for uncatalysed reaction activation energy required.
This figure represents the effect
of an enzyme on an exergonic
activation energy reaction. The activation energy
for catalysed reaction
is needed to destabilize
the chemical bonds in the
reactant reactant. The upper curve
shows the activation energy
energy released

when no enzyme is involved.

The lower curve shows the
activation energy required
when an enzyme is present to
catalyse the reaction.

Mechanism of enzyme action

The following summarizes the mechanism of enzyme action.
The surface of the substrate contacts the active site of the enzyme.
The enzyme changes shape to accommodate the substrate.
A temporary complex called the enzymesubstrate complex forms.
The activation energy is lowered and the substrate is altered by the rearrangement of
the existing atoms.
The transformed substrate, the product, is released from the active site.
The unchanged enzyme is then free to combine with other substrate molecules.
Enzyme action can also be summarized by the following equation:
E + S ES E + P
where E is the enzyme, S is the substrate, ES is the enzymesubstrate complex, and P is
the product.

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08 Metabolism, cell respiration, and photosynthesis

A coenzyme is not Inhibition

usually a protein. It has The effects of pH, temperature, and substrate concentration on the action of enzymes
an essential role in the
normal actions of an were discussed in Section 2.5. Here, we will discuss the effects of certain types of
enzyme. molecules on enzyme active sites. If a molecule affects an active site in some way, the
activity of the enzyme may be altered.

Competitive inhibition
In competitive inhibition, a molecule called a competitive inhibitor competes directly
substrate with the usual substrate for the active site of an enzyme. The result is that the substrate
will have fewer encounters with the active site and rate of the chemical reaction will
competitive be decreased. The competitive inhibitor must have a structure similar to the substrate
to function in this way. An example is the use of sulfanilamide (a sulfa drug) to kill
bacteria during an infection. Folic acid is essential to bacteria as a coenzyme. It is
produced in bacterial cells by enzyme action on paraminobenzoic acid (PABA). The
sulfanilamide competes with the PABA and blocks the enzyme. This prevents the
production of folic acid resulting in the death of the bacteria. Because human cells do
not use PABA to produce folic acid, they are unaffected by the drug.
Figure 8.3 A competitive Competitive inhibition may be reversible or irreversible. Reversible competitive
inhibitor blocks the active site
inhibition may be overcome by increasing the substrate concentration. By doing
of an enzyme so the substrate
cannot bind to it. this, there are more substrate molecules to bind with the active sites as they become
available, and the chemical reaction may proceed more rapidly.

substrate Non-competitive inhibition

Non-competitive inhibition involves an inhibitor that does not compete for the
enzymes active site. In this case, the inhibitor interacts with another site on the
enzyme (see Figure 8.4). Non-competitive inhibition is also referred to as allosteric
inhibition, and the site the inhibitor binds to is called the allosteric site. Binding at
enzyme the allosteric site causes a change in the shape of the enzymes active site, making it
non-functional. Examples of non-competitive inhibition include metallic ions, such
as mercury, binding to the sulfur groups of the component amino acids of many
allosteric inhibitor enzymes. This results in shape changes of the protein, which causes inhibition of the
Figure 8.4 An allosteric (non-
competitive) inhibitor combines Again, this type of inhibition may be reversible or irreversible. There are also examples
with the allosteric site of an of allosteric interactions activating an enzyme rather than inhibiting it.
enzyme, causing the active site
to change shape; the substrate
cannot then bind to it. End-product inhibition
End-product inhibition prevents the cell from wasting chemical resources and energy
by making more of a substance than it needs. Many metabolic reactions occur in an
assembly-line type of process so that a specific end product can be achieved. Each step
of the assembly line is catalysed by a specific enzyme. When the end product is present
in a sufficient quantity, the assembly line is shut down. This is usually done by inhibiting
the action of the enzyme in the first step of the pathway. As the existing end product
is used up by the cell, the first enzyme is reactivated. The enzyme that is inhibited and
reactivated is an allosteric enzyme. When present in higher concentrations, the end
product binds with the allosteric site of the first enzyme, thus bringing about inhibition.
Lower concentrations of the end product result in fewer bindings with the allosteric site
of the first enzyme, and, therefore, activation of the enzyme.

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initial initial
substrate substrate

Figure 8.5 A short pathway

enzyme 1  enzyme 1 of metabolic reactions with
a specific end product that,
when in sufficient quantity,
causes end-product inhibition.
intermediate intermediate This is also a form of negative
A A feedback. The intermediates
are essential molecules
produced in the step-by-
enzyme 2 enzyme 2 step pathway to achieve the
end product. Arepresents
a normal pathway with
several enzymes producing
intermediate intermediate intermediate compounds
B B along the way. B represents
feedback inhibition. In this
condition a large amount of
enzyme 3 enzyme 3 end-product is present. The
end-product inhibits enzyme
1 in the pathway. The result is
that the pathway is halted.
end product end product

The bacterium Escherichia coli uses a metabolic pathway to produce the amino acid
isoleucine from threonine. It is a five-step process. If isoleucine is added to the growth
medium of E. coli, it inhibits the first enzyme in the pathway and isoleucine will not be
synthesized. This situation will exist until the isoleucine is used up.
The inhibition of the first enzyme in the pathway prevents the build-up of
intermediates in the cell. This is a form of negative feedback.

Many examples of enzyme inhibitors exist in medicine. Two examples of competitive inhibitors
are ethanol and fomepizole. Either one of these two may be used as an antidote for ethylene
glycol or methanol poisoning. Ethylene glycol and methanol may be used in producing car
antifreeze, de-icing solutions, solvents, and cleaners. Fomepizole is a competitive inhibitor of
alcohol dehydrogenase, which catalyses the breakdown of ethylene glycol and methanol into
toxic metabolites. This allows other catalytic pathways to be activated, which do not result in
toxic substances.

Worked example
Competitive and non-competitive inhibitors are examples of reversible enzyme
inhibitors. When graphs of their effects are produced, certain characteristics can be
seen. When a chemical is a competitive inhibitor, it competes for the active site of
an enzyme, and its concentration must be kept high to keep the chemical reaction
occurring at a slower rate. Non-competitive inhibitors do not compete for the active
site of the enzyme. The result of this is that the rate of reaction will only increase if
the enzyme concentration is increased. Examine Figure 8.6.

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08 Metabolism, cell respiration, and photosynthesis


Figure 8.6 Enzyme inhibition. V C
[S] = substrate concentration;
V = reaction rate; Vmax =
maximum reaction rate.


Curve A represents a chemical reaction catalysed by an enzyme without the effect

of an inhibitor. Curves B and C represent chemical reactions catalysed by enzymes
affected by inhibitors.
1 Which curve represents the reaction in which a competitive inhibitor is active?
2 Which curve represents the effects of a non-competitive inhibitor?
3 Explain your answers.

1 Curve B.
When asked to
2 Curve C.
differentiate between
competitive and non- 3 Curve C shows the action of a non-competitive inhibitor because it results
competitive inhibition in a lower maximum reaction rate. This occurs because the inhibitor binds
curves on a graph, look to the enzyme present and is not released. The reaction rate will not increase
to see if Vmax is achieved
as the substrate increases because there is a limited amount of enzymes still
as the substrate is
increased. If it is, then active. Curve B represents competitive inhibition because, as the substrate
competitive inhibition increases, the rate of the reaction also increases. This is because of the larger
is being represented. If concentration of substrate out-competing the inhibitor for the active site of the
it is not achieved and is
enzyme. Curve B will eventually equal the maximum reaction when enough
significantly less, non-
competitive inhibition is substrate is added.
being represented.

Developments in areas such as bioinformatics have enhanced the research into metabolic
pathways. Bioinformatics uses many areas of computer science and mathematics to look for
unique events and patterns. This research uses large amounts of data stored in databases.
Scientists often develop software to help this type of research. In 2011, an international team
from the Genomics Institute of the Novartis Research Foundation and The Scripps Research
Institution, by studying databases, discovered a new group of compounds that may lead to a
new generation of anti-malarial drugs capable of both preventing the disease and of alleviating
symptoms when the disease is already present in an individual.
This discovery came after mining the data for groups of related compounds that showed
activity in the liver. They found a cluster of compounds related to imidazolopiperazine that is
showing great promise.

To learn more about how Exercises

enzymes work, go to the 1 Explain why enzymes only work with specific substrates.
hotlinks site, search for the
2 What determines whether an enzyme is competitively or non-competitively inhibited?
title or ISBN, and click on
Chapter 8: Section 8.1. 3 Where would the most efficient control of a metabolic pathway involving end-product inhibition


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8.2 Cell respiration Paradigm shift: the

chemiosmotic theory led
to a paradigm shift in the
field of bioenergetics.
Cell respiration involves the oxidation and reduction of electron carriers.
Phosphorylation of molecules makes them less stable.
In glycolysis, glucose is converted to pyruvate in the cytoplasm.

Glycolysis gives a small net gain of ATP without the use of oxygen.

In aerobic cell respiration pyruvate is decarboxylated and oxidized, and converted into acetyl

compound and attached to coenzyme A to form acetyl coenzyme A in the link reaction.
In the Krebs cycle, the oxidation of acetyl groups is coupled to the reduction of hydrogen carriers,

liberating carbon dioxide.

Energy released by oxidation reactions is carried to the cristae of the mitochondria by reduced

NAD and FAD.

Transfer of electrons between carriers in the electron transport chain in the membrane of the

cristae is coupled to proton pumping.

In chemiosmosis protons diffuse through ATP synthase to generate ATP.

Oxygen is needed to bind with the free protons to form water to maintain the hydrogen gradient,

resulting in the formation of water.

The structure of the mitochondrion is adapted to the function it performs.

Applications and skills:

Application: Electron tomography used to produce images of active mitochondria.
Skill: Analysis of diagrams of the pathways of aerobic respiration to deduce where decarboxylation
and oxidation reactions occur.
Skill: Annotation of a diagram of a mitochondrion to indicate the adaptations to its functions.

The names of the intermediate compounds in glycolysis and the Krebs cycle are not required.

Oxidation and reduction

In Chapter 2 the general processes of respiration and photosynthesis were discussed.
In this chapter we will consider these aspects of cellular metabolism in detail. It is
important to recall that metabolism is the sum total of all the chemical reactions
carried out by an organism. These reactions involve:
catabolic pathways
anabolic pathways.
Catabolic pathways result in the breakdown of complex molecules into smaller
molecules. Conversely, anabolic pathways result in the synthesis of more complex
molecules from simpler ones. Cellular respiration is an example of a catabolic pathway.
Photosynthesis is an example of an anabolic pathway. To understand these complex
pathways, it is essential to understand two general types of chemical reactions:
oxidation and reduction.


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08 Metabolism, cell respiration, and photosynthesis

Oxidation and reduction can be compared using a table like Table 8.2.

Table 8.2 A comparison of oxidation and reduction

Oxidation Reduction

Loss of electrons Gain of electrons

Gain of oxygen Loss of oxygen

Loss of hydrogen Gain of hydrogen

If asked in an exam
to compare oxidation Results in many CO bonds Results in many CH bonds
and reduction, using a
table like Table 8.2 is an Results in a compound with lower Results in a compound with higher
excellent way to structure potential energy potential energy
the answer.
A useful way to remember the general meaning of oxidation and reduction is to think
of the words oil rig.
oil = oxidation is loss (of electrons)
rig = reduction is gain (of electrons).
These two reactions occur together during chemical reactions. Think of it in this way:
Benedicts reagent is
one compounds or elements loss is another compounds or elements gain. This is
a chemical reagent shown by the following equation:
commonly used to detect
the presence of simple
C6H12O6 + 6 O2 6 CO2 + 6 H2O + energy
reducing sugars. It contains In this equation, glucose is oxidized because electrons are transferred from it to
soluble blue copper (II)
ions that may be reduced
oxygen. The protons follow the electrons to produce water. The oxygen atoms that
to copper (I) ions. These occur in the oxygen molecules on the reactant side of the equation are reduced.
copper (I) ions are not Because of this reaction, there is a large drop in the potential energy of the compounds
soluble in water and on the product side of the equation.
will form a redorange
coloured precipitate. Because oxidation and reduction always occur together, these chemical reactions are
The colour of the referred to as redox reactions. When redox reactions take place, the reduced form of
precipitate indicates the
quantity of simple sugar
a molecule always has more potential energy that the oxidized form of the molecule.
present: a green colour Redox reactions play a key role in the flow of energy through living systems. This is
indicates a low sugar because the electrons that are flowing from one molecule to the next are carrying
concentration and a red energy with them. In a similar sort of way, the catabolic and anabolic pathways
colour indicates a high
mentioned earlier are also closely associated with one another. You will see this
sugar concentration. The
electrons to reduce the association as you work through this chapter.
Benedicts reagent resulting
in the colour change come
from the oxidation of the An overview of respiration
sugar molecules. Section 2.8 provided an introduction to the process of cellular respiration. Three
aspects of cellular respiration were discussed:
glycolysis anaerobic respiration aerobic respiration.
As you will recall, glycolysis occurs in the cytoplasm of the cell, produces small
amounts of adenosine triphosphate (ATP) and ends with the product known as
pyruvate. If no oxygen is available, the pyruvate enters into anaerobic respiration. This
occurs in the cytoplasm and it does not result in any further production of ATP. The
products of anaerobic respiration are lactate or ethanol and carbon dioxide. If oxygen
is available, the pyruvate enters aerobic respiration in the mitochondria of the cell. This
process results in the production of a large number of ATPs, carbon dioxide and water.

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In this section, we will discuss the cellular respiration that involves glycolysis and the
three stages of aerobic respiration:
the link reaction
the Krebs cycle
oxidative phosphorylation.

The word glycolysis means sugar splitting and this pathway is thought to be one of
the first biochemical pathways to evolve. It uses no oxygen and occurs in the cytosol of
the cell. No organelles are required. The sugar splitting proceeds efficiently in aerobic
and anaerobic environments. Glycolysis occurs in both prokaryotic and eukaryotic
cells. A hexose, usually glucose, is split in the process. This splitting actually involves
many steps but we can explain it effectively in three stages.
1 Two molecules of ATP are used to begin glycolysis. In the first reaction, the
phosphates from the ATPs are added to glucose to form fructose-1,6-biphosphate,
a process called phosphorylation. The importance of phosphorylation in this step
is that it creates a less stable molecule.

Figure 8.7 The first stage of

6-carbon glucose glycolysis; the circles represent
carbon atoms.



fructose-1, 6-bisphosphate

2 The less stable 6-carbon phosphorylated fructose is split into two 3-carbon sugars
called glyceraldehyde-3-phosphate (G3P). This splitting process is known as lysis.


Figure 8.8 The second stage

of glycolysis.
fructose-1, 6-bisphosphate


glyceraldehyde-3-phosphate glyceraldehyde-3-phosphate


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08 Metabolism, cell respiration, and photosynthesis

3 Once the two G3P molecules are formed, they enter an oxidation phase involving
ATP formation and the production of the reduced coenzyme NAD. Each G3P or
triose phosphate molecule undergoes oxidation to form a reduced molecule of
NAD+, which is NADH. As NADH is being formed, released energy is used to add
an inorganic phosphate to the remaining 3-carbon compound. This results in a
compound with two phosphate groups. Enzymes then remove the phosphate
groups so that they can be added to adenosine diphosphate (ADP) to produce ATP.
The end result is the formation of four molecules of ATP, two molecules of NADP,
and two molecules of pyruvate. Pyruvate is the ionized form of pyruvic acid.

Figure 8.9 The third stage of


2 P i
Once pyruvate is
obtained, the next
pathway is determined P P
by the presence of
oxygen. If oxygen is
present, pyruvate enters
the mitochondria and
aerobic respiration
occurs. If oxygen is
not present, anaerobic
respiration occurs in
the cytoplasm. In the
latter case, pyruvate is 2
converted to lactase in
animals, and ethanol
and carbon dioxide in
Summary of glycolysis
Two ATPs are used to start the process.
The way of producing A total of four ATPs are produced: a net gain of two ATPs.
ATP in glycolysis is
called substrate-level Two molecules of NADH are produced.
phosphorylation because The pathway involves substrate-level phosphorylation, lysis, oxidation, and ATP
the phosphate group is formation.
transferred directly to
ADP from the original The pathway occurs in the cytoplasm of the cell.
phosphate-bearing This metabolic pathway is controlled by enzymes. Whenever ATP levels in the cell are
molecule. high, feedback inhibition will block the first enzyme of the pathway. This will slow or
stop the process.
Two pyruvate molecules are present at the end of the pathway.

To learn more about Mitochondria

mitochondria, go to the It is inside the mitochondria and in the presence of oxygen that the remainder of
hotlinks site, search for the
title or ISBN, and click on cellular respiration occurs.
Chapter 8: Section 8.2. We discussed the structure of the mitochondrion in Chapter 1. You might like to
refresh your memory of this because, as we discuss aerobic respiration, which occurs
in the mitochondrion, we will refer to parts of this organelle.

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This false-colour transmission
electron micrograph (TEM) of
a mitochondrion shows the
internal structure. The matrix
(blue) is permeated by the
membranous cristae (pink).

The link reaction and the Krebs cycle

Once glycolysis has occurred and there is oxygen present, pyruvate enters the matrix
of the mitochondria via active transport. Inside, pyruvate is decarboxylated, a reaction
involving the loss of a carbon in the form of carbon dioxide, to form the 2-carbon
acetyl group. This is the link reaction shown in Figure 8.10. The removed carbon is
released as carbon dioxide, a waste gas. The acetyl group is then oxidized with the
formation of reduced NAD+. Finally, the acetyl group combines with coenzyme A
(CoA) to form acetyl CoA.

NAD coenzyme A S CoA Figure 8.10 The link reaction.
CH3 acetyl CoA Decarboxylation is the
pyruvate transport protein removal of a carbon
Cytosol Mitochondrion


The link reaction is controlled by a system of enzymes. The greatest significance of this A coenzyme is a molecule
reaction is that it produces acetyl CoA. Acetyl CoA may then enter the Krebs cycle to that aids an enzyme in its
action. Coenzymes usually
continue the aerobic respiration process.
act as electron donors or
So far in this discussion, the respiratory substrate has been a hexose. However, in acceptors.
reality, acetyl CoA can be produced from most carbohydrates and fats. Acetyl CoA can
be synthesized into a lipid for storage purposes. This occurs when ATP levels in the cell
are high.
If cellular ATP levels are low, the acetyl CoA enters the Krebs cycle. This cycle is also
called the tricarboxylic acid cycle. It occurs in the matrix of the mitochondrion and
is referred to as a cycle because it begins and ends with the same substance. This is a

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08 Metabolism, cell respiration, and photosynthesis

characteristic of all cyclical pathways in metabolism. You do not need to remember the
names of all the compounds formed in the Krebs cycle. However, it is important that
you understand the overall process.
Lets consider the cycle as a series of steps.
1 Acetyl CoA from the link reaction combines with a 4-carbon compound called
oxaloacetate. The result is a 6-carbon compound called citrate.
Figure 8.11 Acetyl CoA acetyl
combines with oxaloacetate to
form citrate. CoA

oxaloacetate citrate
4C 6C

2 Citrate (a 6-carbon compound) is oxidized to form a 5-carbon compound. In this

process, the carbon is released from the cell (after combining with oxygen) as
carbon dioxide. While the 6-carbon compound is oxidized, NAD+ is reduced to
form NADH.
Figure 8.12 Then a 5-carbon acetyl CoA
compound is formed.

4C 6C CO2



3 The 5-carbon compound is oxidized and decarboxylated to form a 4-carbon

compound. Again, the removed carbon combines with oxygen and is released as
carbon dioxide. Another NAD+ is reduced to form NADH.
Figure 8.13 Next, a 4-carbon acetyl CoA
compound is produced. CoA

4C 6C





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4 The 4-carbon compound undergoes various changes resulting in several products.
One product is another NADH. The coenzyme FAD is reduced to form FADH2.
There is also a reduction of an ADP to form ATP. The 4-carbon compound
is changed during these steps to re-form the starting compound of the cycle,
oxaloacetate. The oxaloacetate may then begin the cycle again.
acetyl CoA
CoA Figure 8.14 Finally, the
4-carbon compound is
converted to oxaloacetate.
oxaloacetate 6C






It is important to remember that the Krebs cycle will run twice for each glucose
molecule entering cellular respiration. This is because a glucose molecule forms two
pyruvate molecules. Each pyruvate produces one acetyl CoA that enters the cycle.
Look again at the complete Krebs cycle (Figure 8.14) and note the following products
that result from the breakdown of one glucose molecule:
2 ATP molecules per molecule of glucose
6 molecules of NADH (which allow energy storage and transfer)
2 molecules of FADH2
4 molecules of carbon dioxide (released).

Two carbon dioxides are released for each glucose molecule during the link reaction.
Four carbon dioxides are released during the Krebs cycle. This accounts for all six of the
carbon atoms that were present in the initial glucose molecule. Glucose is completely
catabolized and its original energy is now carried by NADH and FADH2 or is in ATP.

So far, only four ATPs have been gained, six are generated (four from glycolysis and
two from the Krebs cycle) but two are used to start the process of glycolysis. Each of
these ATPs has been produced by substrate-level phosphorylation.
Ultimately, the breakdown of each glucose molecule results in a net gain of 36 ATPs.
Lets now consider the phase of cellular respiration where most of the ATPs are
produced. In this phase oxidative phosphorylation is the means by which the ATPs are


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08 Metabolism, cell respiration, and photosynthesis

1 Examine Figure 8.15.
acetyl CoA

oxaloacetate 6C


Figure 8.15 NAD





(a) Redraw the figure and place an arrow and the letter D at the two locations where
decarboxylation occurs. On the same figure place an arrow and the letter O at the five
locations where oxidation of a cyclic intermediate compound occurs.
(b) How did you determine the two locations where decarboxylation occurred?
(c) How did you determine the five locations where oxidation occurred?

To learn more about the Electron transport chain and chemiosmosis

electron transport chain, The electron transport chain is where most of the ATPs from glucose catabolism are
go to the hotlinks site,
search for the title or ISBN,
produced. It is the first stage of cellular respiration where oxygen is actually needed,
and click on Chapter 8: and it occurs within the mitochondrion. However, unlike the Krebs cycle, which
Section 8.2. occurred in the matrix, the electron transport chain occurs on the inner mitochondrial
membrane and on the membranes of the cristae.
Embedded in the membranes involved are molecules that are easily reduced and
oxidized. These carriers of electrons (energy) are close together and pass the electrons
The haem group of the from one to another because of an energy gradient. Each carrier molecule has a slightly
carrier is the part that different electronegativity, and, therefore, a different attraction for electrons. Most of
is easily reduced and these carriers are proteins with haem groups and are referred to as cytochromes. One
carrier is not a protein and is called coenzyme Q.
In this chain, electrons pass from one carrier to another because the receiving
molecule has a higher electronegativity and, therefore, a stronger attraction for
electrons. In the process of electron transport, small amounts of energy are released.
The sources of the electrons that move down the electron transport chain are the
coenzymes NADH and FADH2 from the previous stages of cellular respiration.


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40 FeS
free energy relative to O2 / k cal mol1

35 Q
Cyt b

30 FeS

Cyt c1
25 Cyt c
Cyt a
Cyt a3
20 Carriers

FMN: This protein carrier has

15 a flavin-containing group.
FeS: This protein contains an iron
sulfur complex.
Cyt: These are cytochromes (iron- Figure 8.16 The oxidation
containing proteins). reduction reactions of the
5 Q: This is coenzyme Q, also called electron transport chain. It
ubiquinone; it is not a protein. is not necessary for you to
remember all the names of the
1 carriers.
0 2 O2
reductionoxidation reactions

In Figure 8.16 it is clear that the electrons are stepping down in potential energy as they
pass from one carrier to another. It is important to note that: The kangaroo rat from
a desert region of the
FADH2 enters the electron transport chain at a lower free energy level than NADH, USA gets 90% of its daily
thus FADH2 allows the production of two ATPs while NADH allows the production water intake from water of
of three ATPs metabolism. In contrast,
a typical human only
at the very end of the chain, the de-energized electrons combine with available gets 12% of his or her
oxygen. daily water intake from
Oxygen is the final electron acceptor because it has a very high electronegativity and,
therefore, a strong attraction for electrons. When the electrons combine with the
oxygen, so do two hydrogen ions from the aqueous surroundings. The result is water.
Because of the way this water is formed, it is referred to as water of metabolism.
It is also clear from Figure 8.16 that there is a fairly large number of electron carriers. No ATPs are produced
Because of the larger number, the electronegativity difference between adjacent directly by the
electron transport
carriers is not as great. This means that lower amounts of energy are lost at each chain. However, this
exchange. These lower amounts of energy are effectively harnessed by the cell to carry chain is essential to
out phosphorylation. If the amount of energy lost at each exchange was high, much of chemiosmosis, which
it could not be used and the cell could even be damaged. does produce the ATP.


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08 Metabolism, cell respiration, and photosynthesis

So energy is now available as a result of the electron transport chain. This is the energy
that allows the addition of phosphate and energy to ADP to form ATP. The process by
which this occurs is called chemiosmosis. Chemiosmosis involves the movement of
protons (hydrogen ions) to provide energy so that phosphorylation can occur. Because
this type of phosphorylation utilizes an electron transport chain, it is called oxidative
phosphorylation. Substrate-level phosphorylation, mentioned in the earlier phases of
cellular respiration, does not involve an electron transport chain.
Before continuing, it is essential to review the interior structure of the mitochondrion.
In the process of cellular respiration, the structure of the mitochondrion is very
Using any diagram or closely linked to its function. The matrix is the area where the Krebs cycle occurs. The
photomicrograph of a
cristae provide a large surface area for the electron transport chain to function on.
mitochondrion, practise
annotating where the The membranes also provide a barrier, allowing proton accumulation on one side.
processes of aerobic Embedded in the membranes are the enzymes and other compounds necessary for the
respiration occur. processes of the electron transport chain and chemiosmosis to occur.
The inner membranes of the mitochondria have numerous copies of an enzyme
called ATP synthase. This enzyme uses the energy of an ion gradient to allow the
phosphorylation of ADP. The ion gradient is created by a hydrogen ion concentration
difference that occurs across the cristae membranes. Figure 8.17 shows oxidative

intermembrane electron H H H H
H H Cyt c
space carrier
protein complex
of electron
carriers Q

inner 1 synthase
2 H  2 O2 H2O
mitochondrial FADH2 FAD
membrane ATP

mitochondrial NADH  H H
matrix (carrying electrons from food)

Electron transport chain Chemiosmosis

Figure 8.17 Oxidative

phosphorylation occurs at
In Figure 8.17, note the three labelled areas on the left: the intermembrane space, inner
the inner membranes of mitochondrial membrane, and mitochondrial matrix. Also, note that hydrogen ions
the mitochondria of a cell. are being pumped out of the matrix into the intermembrane space. The energy for this
The pumping actions of pumping action is provided by the electrons as they are de-energized moving through
the carriers result in a high
concentration of hydrogen ions
the electron transport chain. This creates the different hydrogen ion concentration on
in the intermembrane space. the two sides of the cristae membranes, mentioned above. With the higher hydrogen
This accumulation allows ion concentration in the intermembrane space, these ions begin to move passively
movement of the hydrogen through a channel in ATP synthase back into the mitochondrial matrix. As the
ions through the enzyme ATP
hydrogen ions move through the ATP synthase channel, the enzyme harnesses the
synthase. The enzyme uses the
energy from the hydrogen flow available energy, thus allowing the phosphorylation of ADP.
to couple phosphate with ADP
to produce ATP.


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Unfortunately, since the beginning of scientific experimentation, there have been instances of Because of the
improper presentation of results. These improper presentations have included improper data hydrophobic region
reporting or even data fabrication. In the scientific community, this misconduct is extremely of the membrane, the
frowned upon. Discuss the possible repercussions to science research as a whole when such hydrogen ions can
misconduct occurs. Continue your discussion to include reasons why scientists sometimes only pass through the
present improper or fabricated data. ATP synthase channel.
Some poisons that
affect metabolism
Summary of ATP production in cellular respiration act by establishing
alternative pathways
We have now described the complete catabolism of one molecule of glucose. The raw through the membrane,
materials were glucose and oxygen. Many enzymes, carriers, and other molecules are thus preventing ATP
involved in the process. The products are carbon dioxide, water, and ATP. The ATPs are production.
essential because they provide the energy by which life is maintained. We can describe
the energy flow in the general process as shown in Figure 8.18.

Figure 8.18

Glucose NADH or FADH2 Electron transport chain Chemiosmosis ATP

To account for the production of ATP in cellular respiration, lets look at the three main
processes, glycolysis, the Krebs cycle, and the electron transport chain, in a table.

Table 8.3 The processes of cellular respiration

Process ATP used ATP Net ATP

produced gain

Glycolysis 2 4 2

Krebs cycle 0 2 2

Electron transport chain and chemiosmosis 0 32 32

Total 2 38 36

Theoretically 36 ATPs are produced by cellular respiration, but in reality the number
is closer to 30. This is thought to be because some hydrogen ions move back to the
matrix without going through the ATP synthase channel. Also, some of the energy
from hydrogen ion movement is used to transport pyruvate into the mitochondria.
The 30 ATPs generated by cellular respiration account for approximately 30% of the
energy present in the chemical bonds of glucose. The remainder of the energy is lost
from the cell as heat.

A final look at respiration and the mitochondrion

Cellular respiration is the process by which ATP is provided to the organism so that it
can live. It is a very complex series of chemical reactions, most of which occur in the
mitochondrion. Lets end our discussion of this essential-to-life process by looking
at a table showing the parts of the mitochondrion and how those parts allow cellular


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08 Metabolism, cell respiration, and photosynthesis

Table 8.4 The role of the mitochondrion in cellular respiration

Feature Role

Outer mitochondrial A membrane that separates the contents of the

membrane mitochondrion from the rest of the cell

Matrix An internal cytosol-like area that contains the enzymes for

the link reaction and the Krebs cycle

Cristae Tubular regions surrounded by membranes that increase

the surface area for oxidative phosphorylation
Inner mitochondrial A membrane that contains the carriers for the electron
YOURSELF membrane transport chain and ATP synthase for chemiosmosis
2 Annotate the diagram of a
Space between inner and A reservoir for hydrogen ions (protons)
mitochondrion provided in
outer membranes
Figure 8.19 to indicate the
adaptations that allow the
mitochondrion to carry out
The overall equation for cellular respiration is:
its essential functions. C6H12O6 + 6 O2 6 CO2 + 6 H2O + energy (as heat or ATP)
All organisms need the ability to produce ATP for energy, so all organisms carry out

4 Using ideal ATP production numbers, how many ATPs would an individual generate if he or she
Figure 8.19 consumed only pyruvate and carried one pyruvate molecule through cellular respiration?
5 Striated or voluntary muscles that occur in humans generally have a larger number of mitochondria
than other cell types. Why is this important?
6 If both NAD and FAD are reduced, which would allow the greater production of ATPs via the electron
transport chain and chemiosmosis?
7 If an individual took a chemical that increased the ability of hydrogen ions to move through the
phospholipid bilayer of the mitochondrial membranes, what would the effect be on ATP production?
8 If ATP synthase was not present in the cristae of a mitochondrion, what would be the effect?

Developments in
scientific research
8.3 Photosynthesis
follow improvements
in apparatus: sources of
C and autoradiography Understandings:
enabled Calvin to Light-dependent reactions take place in the intermembrane space of the thylakoids.
elucidate the pathways of Light-independent reactions take place in the stroma.
carbon fixation. Reduced NADP and ATP are produced in the light-dependent reactions.

Absorption of light by photosystems generates excited electrons.

Photolysis of water generates electrons for use in the light-dependent reactions.

Transfer of excited electrons occurs between carriers in thylakoid membranes.

To see an introduction to
Excited electrons from Photosystem II are used to contribute to generate a proton gradient.
photosynthesis, go to the
ATP synthase in thylakoids generates ATP using the proton gradient.
hotlinks site, search for the
Excited electrons from Photosystem I are used to reduce NADP.
title or ISBN, and click on
In the light-independent reactions a carboxylase catalyses the carboxylation of ribulose
Chapter 8: Section 8.3.
Glycerate 3-phosphate is reduced to triose phosphate using reduced NADP and ATP.

Triose phosphate is used to regenerate RuBP and produce carbohydrates.

Ribulose bisphosphate is reformed using ATP.

The structure of the chloroplast is adapted to its function in photosynthesis.


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Applications and skills:
Application: Calvins experiment to elucidate the carboxylation of RuBP.
Skill: Annotation of a diagram to indicate the adaptations of a chloroplast to its function.

The chloroplast
Some people refer to the chloroplast as a photosynthetic machine. They are not wrong.
Unlike respiration, where some of the steps occur outside the mitochondrion, all of
the photosynthetic process occurs within the chloroplast. Chloroplasts, along with
Figure 8.20 This false-colour
mitochondria, represent possible evidence for the theory of endosymbiosis, discussed
TEM and drawing show the
in Chapter 1, Section 1.5. Both organelles have an extra outer membrane (indicating a structure of a chloroplast. Can
need for protection in a potentially hostile environment), their own DNA, and they are you find as many parts in the
very near in size to a typical prokaryotic cell. TEM as are labelled in the
outer membrane

inner membrane
granum of several

1 thylakoid

The structure of the chloroplast was discussed in Chapter 1. You may want to return
to that chapter for a brief refresher. Chloroplasts occur mostly within the cells of the
photosynthetic factory of the plant, the leaves. However, some plants have chloroplasts
in cells of other organs.

Plastids are a group of closely related organelles that occur in photosynthetic eukaryotic cells.
There are three types of plastid that occur in plant cells:
chloroplasts, which are green and involved in photosynthesis
leucoplasts, which are white or clear and function as energy storehouses
chromoplasts, which are brightly coloured and synthesize and store large amounts of
orange, red, or yellow pigments.
All these plastids develop from a common proplastid.

The overall process of photosynthesis

During the discussion on respiration, we considered the means by which the cell
breaks down chemical bonds in glucose to produce ATP. In this section, the discussion
centres on the establishment of chemical bonds to produce organic compounds. Using
light energy, the raw materials of photosynthesis are carbon dioxide and water. Many
enzymes are involved to enable the formation of products that include glucose, more
water, and oxygen. The overall equation is:
6 CO2 + 12 H2O C6H12O6 + 6 H2O + 6 O2

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08 Metabolism, cell respiration, and photosynthesis

Water occurs on both sides because 12 molecules are consumed and 6 molecules
are produced. Clearly, photosynthesis is essentially the reverse of respiration.
Whereas respiration is, in general, a catabolic process, photosynthesis is, in general,
an anabolic process. Photosynthesis occurs in organisms referred to as autotrophs.
These organisms make their own food. Non-photosynthetic and non-chemosynthetic
organisms are referred to as heterotrophs. They must obtain their food (which is
necessary for energy) from other organisms.
Photosynthesis involves two major stages:
the light-dependent reaction
Light energy behaves as if
it exists in discrete packets the light-independent reaction.
called photons. Shorter
wavelengths of light have The light-dependent reaction
greater energy within
their photons than longer The light-dependent reaction occurs in the thylakoids or grana of the chloroplast. A
wavelengths. Photons can stack of thylakoids make up a granum (plural grana). Light supplies the energy for
transfer their energy upon this reaction to occur. The ultimate source of light is the Sun. Even though plants may
interaction with other
survive quite well when they receive light from sources other than the Sun, most plants
particles. This transfer of
energy occurs many times on our planet rely on the Sun for the energy necessary to drive photosynthesis.
in photosynthesis.
To absorb light, plants have special molecules called pigments. There are several
different pigments in plants, and each effectively absorbs photons of light at different
wavelengths. The two major groups are the chlorophylls and the carotenoids.
These pigments are organized on the membranes of the thylakoids. The regions of
organization are called photosystems and include:
chlorophyll a molecules
accessory pigments
a protein matrix.
The reaction centre is the portion of the photosystem that contains:
a pair of chlorophyll a molecules
a matrix of protein
a primary electron acceptor.
Bacteria that carry out photosynthesis have only one type of photosystem. However,
modern-day plants have two types of photosystem. Each absorbs light most efficiently
at a different wavelength. Photosystem I is most efficient at 700 nanometres (nm) and
is labelled as P700. Photosystem II is most efficient at 680 nm and is labelled as P680.
These two photosystems work together to bring about a non-cyclical electron transfer.
Figure 8.21 shows the overall light-dependent reaction of photosynthesis involving
non-cyclic photophosphorylation (non-cyclic electron flow).
The numbered descriptions that follow refer to the numbered steps in Figure 8.21.
1 A photon of light is absorbed by a pigment in Photosystem II and is transferred
to other pigment molecules until it reaches one of the chlorophyll a (P680)
molecules in the reaction centre. The photon energy excites one of the chlorophyll
a electrons to a higher energy state.
2 This electron is captured by the primary acceptor of the reaction centre.
3 Water is split by an enzyme to produce electrons, hydrogen ions, and an oxygen
atom. This process is driven by the energy from light and is called photolysis. The
electrons are supplied one by one to the chlorophyll a molecules of the reaction

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electron ferredoxin (electron carrier)
excited e
electron NADP
acceptor reductase
electron carrier (PQ)
complex electron carrier
(electron carrier)
energy of electrons

reaction photon

photon (P680)
reaction water-splitting
centre enzyme

H2 O

2H  O2

Photosystem II (P680) Photosystem I (P700)

Figure 8.21 The light-

dependent reaction of
4 The excited electrons pass from the primary acceptor down an electron
transport chain, losing energy at each exchange. The first of the three carriers is
plastoquinone (PQ). The middle carrier is a cytochrome complex.
5 The energy lost from the electrons moving down the electron transport
chain drives chemiosmosis (similar to that in respiration) to bring about
phosphorylation of ADP to produce ATP.
6 A photon of light is absorbed by a pigment in Photosystem I. This energy is
transferred through several accessory pigments until received by a chlorophyll
a (P700) molecule. This results in an electron with a higher energy state being
transferred to the primary electron acceptor. The de-energized electron from
Photosystem II fills the void left by the newly energized electron.
7 The electron with the higher energy state is then passed down a second electron
transport chain that involves the carrier ferredoxin.
8 The enzyme NADP reductase catalyses the transfer of the electron from ferredoxin
to the energy carrier NADP+. Two electrons are required to reduce NADP+ fully to
NADPH and ATP are the final products of the light-dependent reaction. They supply
chemical energy for the light-independent reaction to occur. The explanation above
also shows the origin of the oxygen released by photosynthesizing plants (step 3).
However, you need to know more detail about the production of ATP.
ATP production in photosynthesis is very similar to ATP production in respiration.
Chemiosmosis allows the process of phosphorylation of ADP. In this case, the energy
to drive chemiosmosis comes from light. As a result, we refer to the production of ATP
in photosynthesis as photophosphorylation.

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08 Metabolism, cell respiration, and photosynthesis

Besides the non-cyclic A comparison of chemiosmosis in respiration and photosynthesis is shown in

electron pathway used Table 8.5.
to produce ATP by
photophosphorylation, Table 8.5 A comparison of chemiosmosis
there is an alternative
pathway involving Respiration chemiosmosis Photosynthesis chemiosmosis
a cyclic pathway.
This cyclic pathway Involves an electron transport chain Involves an electron transport chain
is discussed on the embedded in the membranes of the embedded in the membranes of the
following page. cristae thylakoids

Energy is released when electrons are Energy is released when electrons are
exchanged from one carrier to another exchanged from one carrier to another

Released energy is used to pump Released energy is used to pump hydrogen

hydrogen ions actively into the ions actively into the thylakoid space
intermembrane space

Hydrogen ions come from the matrix Hydrogen ions come from the stroma

Hydrogen ions diffuse back into the Hydrogen ions diffuse back into the stroma
matrix through the channels of ATP through the channels of ATP synthase

ATP synthase catalyses the ATP synthase catalyses the

phosphorylation of ADP to form ATP photophosphorylation of ADP to form ATP

In both cases ATP synthase is embedded along with the carriers of the electron
transport chain in the membranes involved.
In photosynthesis, the production of ATP occurs between Photosystem II and
Photosystem I. Study Figure 8.22. Notice that the b6f complex, which is a cytochrome
complex, pumps the hydrogen ions into the thylakoid space. This increases the
concentration of these ions, which then move passively through the ATP synthase
channel, providing the energy to phosphorylate ADP.
photon H


PQ membrane
Figure 8.22 Chemiosmosis in
a plant cell chloroplast. H2O H H
Thylakoid H H
1O 2 H H H H H
space 2 2

Photosystem II b6f complex ATP synthase


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The process just described is also known as non-cyclic photophosphorylation. There is another
way the light-dependent reaction of photosynthesis may produce ATP. It is called cyclic
photophosphorylation. It proceeds only when light is not a limiting factor and when there
is an accumulation of NADPH in the chloroplast. In this process, light-energized electrons
from Photosystem I flow back to the cytochrome complex of the electron transport chain
between Photosystem II and Photosystem I (see Figure 8.21). From the cytochrome complex,
the electrons move down the remaining electron transport chain allowing ATP production
via chemiosmosis. These ATPs are then shuttled to the Calvin cycle so that it can proceed
more rapidly.

Experiment to demonstrate electron transfer in chloroplasts

*Safety alerts: Use safety goggles and lab aprons. Be cautious in the use of chemicals and
glassware. Be careful of light sources as they may be hot. Dispose of chemicals as directed by
your teacher. Wash your hands thoroughly upon completion of the activity.*
Electrons energized by light allow the production of ATP and NADPH in the light-dependent
reaction of photosynthesis. In this experiment, DCPIP or DPIP (2,6-dichlorophenol-
indophenol) will be used to replace NADP in the light-dependent reaction. DCPIP is a blue
colour, but turns colourless when reduced. There is a direct relationship between the rate of
photosynthesis and the change in colour from blue DCPIP to its reduced, clear form.
Full details with a worksheet of how to carry out this experiment are available on your eBook.
Data and observations
Table 8.6

Experimental Start colour/ Finish colour/absorbance

cuvette absorbance




1 What was the control in this experiment?
2 What product would have received the energized electrons if DCPIP had not been added?
3 What is the actual source of the electrons that reduced the DCPIP?
4 What was the effect of darkness on the reduction of DCPIP? Explain.
5 What was the effect of boiling the chloroplasts on this experiment? Explain.

The light-independent reaction

The light-independent reaction occurs within the stroma or cytosol-like region of the
The ATP and NADPH produced by the light-dependent reaction provide the energy
and reducing power for the light-independent reaction to occur. Up to this point there
has been no mention of carbohydrate production. Therefore, as we know glucose is
a product of photosynthesis, the result of the light-independent reaction must be the
production of glucose.
The light-independent reaction involves the Calvin cycle (see Figure 8.23), which
occurs in the stroma of the chloroplast. Because it is a cycle, it begins and ends with the
same substance. You should recall that a similar cyclic metabolic pathway occurred in
respiration: the Krebs cycle.

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08 Metabolism, cell respiration, and photosynthesis

6 CO2 from the air

carbon dioxide fixation

6 6
RuBP unstable intermediate

6 ADP 12 ATP


4 P i 12 ADP
12 P i
10 12 NADP
phosphate 12
Figure 8.23 The Calvin cycle. 2 
The numbered steps are triose
described in the text. phosphate
P i

P complex carbohydrates
sugar phosphate

Refer to Figure 8.23 as you read about the steps of the Calvin cycle.
1 Ribulose bisphosphate (RuBP), a 5-carbon compound, binds to an incoming
carbon dioxide molecule in a process called carbon fixation. This fixation is
catalysed by an enzyme called RuBP carboxylase (rubisco). The result is an
unstable 6-carbon compound.
2 The unstable 6-carbon compound breaks down into two 3-carbon compounds
called glycerate 3-phosphate (GP).
3 The 3-carbon molecules of GP are acted upon by ATP and NADPH from the light-
dependent reaction to form two other 3-carbon molecules called triose phosphate
(TP). This is a reduction reaction.
4 The molecules of TP may then go in either of two directions. Some leave the cycle
to become sugar phosphates that may become more complex carbohydrates.
Most, however, continue in the cycle to reproduce the originating compound of
the cycle, RuBP.
5 In order to regain RuBP molecules from TP, the cycle uses ATP.
In Figure 8.23, spheres are used to represent the carbon atoms so that they can be
tracked through the cycle. The coefficients (numbers) in front of each compound
involved show what it takes to produce one molecule of a 6-carbon sugar. It is
clear that for every 12 TP molecules, the cycle produces one 6-carbon sugar and six
molecules of the 5-carbon compound RuBP. All the carbons are accounted for, and the
law of conservation of mass is demonstrated. Also, it is important to note that 18 ATPs

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and 12 NADPH are necessary to produce six RuBP molecules and one molecule of a A team led by Melvin
6-carbon sugar. Calvin in the late 1940s
and early 1950s worked
TP is the pivotal compound in the Calvin cycle. It may be used to produce simple on experiments to find
sugars such as glucose, disaccharides such as sucrose, or polysaccharides such as the early products of
cellulose or starch. However, most of it is used to regain the starting compound of the photosynthesis. The final
Calvin cycle, RuBP. products were already
well known. His research
involved the creation of
NATURE OF SCIENCE the now famous lollipop
apparatus. This specially
As mentioned earlier, Calvin and his team worked out the details of carbon fixation. In order to designed apparatus
do this, Calvin used improvements in apparatus design and recent developments in radioactive was actually a flattened
tracers and autoradiography. Calvin devised the lollipop apparatus. This is a thin, almost bulb- flask that was used to
shaped, glass vessel with a supporting stem. The vessel was designed to mimic the shape of a house algal cells carrying
leaf: thin and broad. He then carried out the following procedures. out photosynthesis. By
Chlorella (a type of green algae) was placed inside the lollipop. using radioactive tracer
The algae cells were then exposed to 14C (radioactive carbon) and light. experiments with this
apparatus, Calvin was
Samples of the Chlorella were then released from the apparatus at short time intervals.
successful in his studies.
Each removed sample was immediately placed into a boiling methanol solution to denature Discuss the role of
the enzymes and stop the photosynthetic process. creativity in scientific
The compounds within the algae were then separated. Two-way paper chromatography investigations as well as
was used for this separation. This process used one solvent to separate the first set of in art.
components. Then the paper was turned and placed in a different solvent to obtain a further
separation of components.
The final radioactive products were identified using autoradiography.
Because Calvin carried out this procedure with algae released at different time intervals, he
obtained different products at different times. This allowed him to sequence the steps of the
overall process and to elucidate the pathways of carbon fixation (the Calvin cycle).

Summary of photosynthesis To learn more about

In summary, the process of photosynthesis includes the light-dependent and the light- photosynthesis, go to the
hotlinks site, search for the
independent reactions. The products of the light-dependent reaction are ATP and
title or ISBN, and click on
NADPH, which are needed to allow the light-independent reaction to proceed. Thus it Chapter 8: Section 8.3
is clear that light is needed for the light-independent reaction to occur, but not directly.
A summary of the two reactions is shown in Figure 8.24.

Light-independent reaction
Light-dependent reactions (Calvin cycle)
ADP Figure 8.24 A summary of
 P i the complete process of
photosystem II RuBP GP
electron transport chain
photosystem I
thylakoid starch
chloroplast cellulose

O2 glucose


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08 Metabolism, cell respiration, and photosynthesis

Note that NADP+ and ATP move back and forth in the chloroplast from the thylakoids
to the stroma in their reduced and oxidized forms. A final summary of the two
reactions is shown in Table 8.7.

Table 8.7 Photosynthesis

Light-dependent reaction Light-independent reaction

Occurs in the thylakoids Occurs in the stroma

Uses light energy to form ATP and NADPH Uses ATP and NADPH to form triose

Splits water in photolysis to provide Returns ADP, inorganic phosphate,

replacement electrons and H+, and to release and NADP to the light-dependent
oxygen to the atmosphere reaction

Includes two electron transport chains and Involves the Calvin cycle
Photosystems I and II

The chloroplast and photosynthesis

From the explanation of photosynthesis, it is clear how important the chloroplast
is to the overall process. The structure of the chloroplast allows the light-dependent
and light-independent reactions to proceed efficiently. In biology, the relationship of
structure to function is a universal theme. The chloroplast and photosynthesis are no
exception to this, as shown in Table 8.8.

Table 8.8 The structure and Chloroplast structure Function allowed

function of a chloroplast
Extensive membrane surface area Allows greater absorption of light by photosystems
of the thylakoids

Small space (lumen) within the Allows for faster accumulation of protons to create
thylakoids a concentration gradient

Stroma region similar to the Allows an area for the enzymes necessary for the
cytosol of the cell Calvin cycle to work in

Double membrane on the Isolates the working parts and enzymes of the
outside chloroplast from the surrounding cytosol


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Scientists in laboratories
CHALLENGE YOURSELF located around the world
are presently working
3 Examine the diagram of a typical chloroplast (Figure 8.25). Answer the questions below the diagram
on the development
with the appropriate letter.
A of an artificial leaf. This
Figure 8.25 leaf would be capable
of carrying out
photosynthesis. The efforts
B of these research facilities
may one day greatly
increase the availability
of food resources for the
worlds hungry human

(a) Which letter represents the stroma where all the enzymes necessary for the light-independent
reaction occur?
To become more
(b) Which letter represents the double membrane that controls the entry and exit of materials for
confident in your
the chloroplast?
understanding of the
(c) What is the letter of the thylakoid that contains the photosystems?
chloroplast, obtain some
(d) Which letter represents a granum, which is where the light-dependent reaction occurs?
electron micrographs of
(e) Which two letters represent the areas of the chloroplasts that cause the green colour of
chloroplasts from several
chloroplasts? Why do these areas create this colour?
different plants. On these
(f) The chloroplasts within some plant cells can often be seen moving in a cyclical pattern near the
micrographs, annotate
periphery of the cell. This is called cyclosis or cytoplasmic streaming. What might be the value
names of structures and
of such movement to the process of photosynthesis?
their functions.

Once you understand the details of photosynthesis, return to Section 2.8. Look again
at the section about limiting factors of photosynthesis. You should now be able to
explain more fully how temperature, light intensity, and carbon dioxide concentration
may limit the rate of photosynthesis.

9 Why do plants need both mitochondria and chloroplasts?
10 You have a leaf from each of two very different plants. One leaf has more pigments than the other.
Which leaf would have the greater photosynthetic rate, assuming all affecting factors are equal? Why?
11 Explain the final products of the two photosystems involved in the light-dependent reaction of
12 Many scientists state that the enzyme RuBP carboxylase (rubisco) is the most ubiquitous protein on
Earth. Why is there a very good chance that this is true?
13 How are the products of the light-dependent reaction important to the light-independent reaction?

The first international conference dedicated to the creation of an artificial leaf was held in
2011. This conference addressed the goals of the Global Artificial Photosynthesis (GAP) project.
Research centres are active in this area of study throughout the world, and the objective of
this conference was to allow the researchers to share their discoveries. Energy capture, energy
conversion and storage, and carbon fixation using modified and synthetic biological processes,
were all addressed. From the GAP project, it is hoped that enhanced crop production, reduced
atmospheric CO2 levels, and increased availability of fuels for heating and cooking may
be realized.


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08 Metabolism, cell respiration, and photosynthesis

Practice questions
1 Where is carbon dioxide produced in the mitochondrion?

(Total 1 mark)
2 In the mitochondrial electron transport chain, what is the last electron acceptor?
A CO2 C O2
(Total 1 mark)
3 Which of the following statements is true about enzymes?
A They are used up in the reactions they catalyse.
B Allosteric inhibitors bind to the active site.
C They lower the energy of activation for a reaction.
D They supply the energy of activation for a reaction.
(Total 1 mark)
4 What is the role of NADH + H in aerobic cell respiration?

A To transfer hydrogen to the electron transport chain.

B To reduce intermediates in the Krebs cycle.
C To accept electrons from the electron transport chain.
D To combine with oxygen to produce water.
(Total 1 mark)
5 What reaction, involving glycerate 3-phosphate, is part of the light-independent reactions of
A Glycerate 3-phosphate is carboxylated using carbon dioxide.
B Two glycerate 3-phosphates are linked together to form one hexose phosphate.
C Glycerate 3-phosphate is reduced to triose phosphate.
D Five glycerate 3-phosphates are converted to three ribulose 5-phosphates.
(Total 1 mark)
6 What is the advantage of having a small volume inside the thylakoids of the chloroplast?
A High proton concentrations are rapidly developed.
B High electron concentrations are rapidly developed.
C Photosynthetic pigments are highly concentrated.
D Enzymes of the Calvin cycle are highly concentrated.
(Total 1 mark)


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7 During glycolysis a hexose sugar is broken down to two pyruvate molecules. What is the correct
sequence of stages?
A Phosphorylation oxidation lysis
B Oxidation phosphorylation lysis
C Phosphorylation lysis oxidation
D Lysis oxidation phosphorylation
Total 1 mark)
8 Which is correct for the non-competitive inhibition of enzymes?

Inhibitor resembles Inhibitor binds to

substrate active site

A yes yes

B yes no

C no yes

D no no
Total 1 mark)
9 Where are the light-dependent and light-independent reactions taking place in the diagram



Light dependent Light independent




Total 1 mark)
10 What is the link reaction in aerobic respiration?
A Pyruvate is carboxylated, acetyl reacts with coenzyme A, reducing NADH + H+
B Pyruvate is decarboxylated, acetyl reacts with coenzyme A, forming NADH + H+
C Pyruvate reacts with coenzyme A, forming NADH + H+
D Pyruvate is decarboxylated, reacting with coenzyme A, reducing NADH + H+
Total 1 mark)


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