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Chapter 1 - Photosynthesis and ATP Synthesis PDF
Chapter 1 - Photosynthesis and ATP Synthesis PDF
Humans, like all animals and fungi, are trapped energy from these molecules and convert
heterotrophs. This means that we need to eat it into a form that their cells can use. This process
food containing organic molecules, especially is called respiration, and it involves oxidation of
carbohydrates, fats and proteins. These organic the energy-containing organic substances, forming
molecules are our only source of energy. another energy-containing substance called ATP.
Plants, however, do not need to take in any Every cell has to make its own ATP. You can find
organic molecules at all. They obtain their energy out more about ATP in Chapter 2.
from sunlight. They can use this energy to build
their own organic molecules for themselves, An overview of photosynthesis
using simple inorganic substances. They first Photosynthesis happens in several different
produce carbohydrates from carbon dioxide and kinds of organisms, not only plants. There are
water, by photosynthesis. They can then use these many kinds of bacteria that can photosynthesise.
carbohydrates, plus inorganic ions such as nitrate, Photosynthesis also takes place in phytoplankton,
phosphate and magnesium, to manufacture all the tiny organisms that float in the upper layers of the
organic molecules that they need. Organisms that sea and lakes. Here, though, we will concentrate
feed in this way self-sufficient, not needing any on photosynthesis in green plants, which takes
organic molecules that another organism has made place in the chloroplasts of several plant tissues,
are autotrophs. especially the palisade mesophyll and spongy
So heterotrophs depend on autotrophs for mesophyll tissues of leaves (Figure 1.1). This
the supply of organic molecules on which they photosynthesis is the ultimate source of almost all
feed. Some heterotrophs feed directly on plants, of our food.
while others feed further along a food chain. But The overall equation for photosynthesis is:
eventually all of an animals or funguss food
6CO2 + 6H2O C6H12O6 + 6O2
can be traced back to plants, and the energy of
sunlight. The xylem tissues of roots, stems andleaf
In this chapter, we will look in detail at how vascular bundles bring water to the photosynthesising
plants transfer energy from sunlight to chemical cells of the leaf. The carbon dioxide diffuses into
energy in organic molecules. In Chapter 2, we will the leaf through stomata, the tiny holes usually
see how all living organisms can then release the found in the lower epidermis of the leaf. It then
1
Chapter 1: Photosynthesis and ATP synthesis
palisade spongy
midrib mesophyll mesophyll
lamina
vascular bundle
cuticle
upper
epidermis
stoma
palisade
mesophyll
veins spongy
lower mesophyll
epidermis
lower
epidermis
2
Chapter 1: Photosynthesis and ATP synthesis
a b
cuticle
upper
epidermis
palisade
mesophyll
cytoplasm
vascular vacuole mesophyll
bundle cell
nucleus
(vein)
chloropast
spongy
mesophyll
air space
lower
epidermis
guard
stoma
cell
conditions. The cuticle and epidermis together turgidity of these guard cells cause them to change
form a protective layer against microorganisms shape so that they open and close the pore. When
and some insects. the guard cells gain water, the pore opens; as they
The structure of the lower epidermis is similar lose water it closes. Guard cells have unevenly
to that of the upper, except that most mesophytes thickened cell walls. The wall adjacent to the pore
have many stomata in the lower epidermis. (Some is very thick, whilst the wall furthest from the
have a few stomata in the upper epidermis also.) pore is thin. Bundles of cellulose microfibrils are
Stomata are the pores in the epidermis through arranged as hoops around each guard cell and,
which diffusion of gases occurs, including carbon as the cell becomes turgid, these hoops ensure
dioxide. Each stoma is bounded by two sausage- that the cell mostly increases in length and not
shaped guard cells (Figure 1.3). Changes in the diameter. Since the ends of the two guard cells are
joined and the thin outer wall bends more readily
than the thick inner one, the guard cells become
stoma
closed curved. This makes the pore between the cells
stoma
open open.
Guard cells gain and lose water by osmosis. A
chloroplasts decrease in water potential is needed before water
can enter the cells by osmosis. This is achieved by
the active removal of hydrogen ions, using energy
guard from ATP, and then intake of potassium ions
cells
(indirect active transport).
An electron micrograph and a drawing of
thick thin
cell wall
a palisade cell is shown in Unit 1 on page 41.
cell wall
Figure 1.4 shows a photomicrograph of palisade
cells. The palisade mesophyll is the main site of
Figure 1.3 Photomicrograph of stomata and photosynthesis, as there are more chloroplasts per
guard cells in Tradescantia leaf epidermis (2000). cell than in the spongy mesophyll.
3
Chapter 1: Photosynthesis and ATP synthesis
large surface area of contact between cell and H2O light-dependent stage O2
air.
The cell walls are thin, so that gases can diffuse
through them more easily.
CO2 light-independent stage
thylakoid SAQ
stroma granum 1 List the features of a chloroplast that aid
Electron micrograph of part of a chloroplast photosynthesis.
lamellae granum stroma thylakoid
ribosome
Photosynthetic pigments
lipid A pigment is a substance whose molecules
droplet
absorb some wavelengths (colours) of light, but
not others. The wavelengths it does not absorb
are either reflected or transmitted through the
substance. These unabsorbed wavelengths reach
(36500)
our eyes, so we see the pigment in these colours.
The majority of the pigments in a chloroplast
Figure 1.6 The structure of a chloroplast. are chlorophyll a and chlorophyll b (Figure 1.7).
5
Chapter 1: Photosynthesis and ATP synthesis
Light energy is
Key absorbed by
chlorophyll a chlorophyll a
chlorophyll b molecules at the
carotene reaction centre.
light
energy The energy is
Light absorbed
passed from
one molecule
to another. Chlorophyll
emits a
high-energy
electron.
light, including more blue-green light than the A low-energy electron a photosystem
replaces the high- including hundreds of
chlorophylls. Theyare accessory pigments. They energy electron that molecules of chlorophyll
help by absorbing wavelengths of light that would was passed on. a, chlorophyll b and
otherwise notbe used by the plant. They pass on carotenoids
photons of light that hit the photosystem, and leaves the chlorophyll molecules completely. The
is funnelled down to a pair of molecules at the electron is then passed along the chain of electron
reaction centre of the photosystem complex. carriers. The energy from the electron is used to
There are two different sorts of photosystem, make ATP. The electron, now having lost its extra
PSI and PSII, both with a small number of energy, eventually returns to chlorophyll a in PSI.
molecules of chlorophyll a at the reaction centre.
Non-cyclic photophosphorylation
Photophosphorylation This process involves both kinds of photosystem.
Photophosphorylation means phosphorylation It results not only in the production of ATP, but
using light. It refers to the production of ATP, by also of reduced NADP.
combining a phosphate group with ADP, using Light hitting either PSI or PSII causes electrons
energy that originally came from light: to be emitted. The electrons from PSII pass down
ADP + phosphate ATP the electron carrier chain, generating ATP by
Photophosphorylation happens when an electron photophosphorylation. However, instead of going
is passed along a series of electron carriers, back to PSII, the electrons instead replace the
forming an electron transport chain in the thylakoid electrons lost from PSI.
membranes. The electron starts off with a lot of The phosphorylation of ADP to ATP involves
energy, and it gradually loses some of it as it moves the movement of H+ across the thylakoid
from one carrier to the next. The energy is used to membrane. This process also occurs in respiration
cause a phosphate group to react with ADP. and is described in detail in Chapter 2.
The electrons emitted from PSI are not used to
Cyclic photophosphorylation make ATP. Instead, they help to reduce NADP.
This process involves only PSI, not PSII. It results For this to happen, hydrogen ions are required.
in the formation of ATP, but not reduced NADP These come from another event that happens when
(Figure 1.10). light hits PSII. PSII contains an enzyme that splits
Light is absorbed by PSI and the energy passed water when it is activated by light. The reaction is
on to electrons in the chlorophyll a molecules at called photolysis:
the reaction centre. In each chlorophyll a molecule, 2H2O 4H+ + 4e + O2
one of the electrons becomes so energetic that it The hydrogen ions are taken up by NADP, forming
reduced NADP. The electrons replace the ones
that were emitted from PSII when light hit it.
high-energy electron The oxygen diffuses out of the chloroplast and
ADP + Pi eventually out of the leaf, as an excretory product.
e
The Z-scheme
ATP
The Z-scheme is simply a way of summarising
photosystem I
high-energy
high-energy e electron
electron
chain of
ADP + Pi
e electron
photosystem I
carriers
energy level
ATP
oxidised NADP + H+
photosystem II
light
absorbed e
reduced NADP
chain of
electron
H2O e
carriers
light
absorbed Key
e change in energy movement of electrons
O2 H
+
water replace the ones that are emitted from the The light-independent stage
photosystem.) Now the ATP and reduced NADP that have been
If you keep following the vertical line showing formed in the light-dependent stage are used
the increasing energy in the electrons, you arrive to help to produce carbohydrates from carbon
at a point where it starts a steep dive downwards. dioxide. These events take place in the stroma
This shows the electrons losing their energy as they of the chloroplast. The cyclic series of reactions is
pass along the electron carrier chain. Eventually known as the Calvin cycle (Figure 1.12).
they arrive at photosystem I. The chloroplast stroma contains an enzyme
You can then track the movement of the called rubisco (its full name is ribulose
electrons to a higher energy level when PSI is hit bisphosphate carboxylase). This is thought to
by light, before they fall back downwards as they be the most abundant enzyme in the world. Its
lose energy and become part of a reduced NADP function is to catalyse the reaction in which carbon
molecule. dioxide combines with a substance called RuBP
SAQ
3 Copy and complete the table to compare cyclic (If a box in a particular row is not applicable,
and non-cyclic photophosphorylation. write n/a.)
Cyclic Non-cyclic
photophosphorylation photophosphorylation
Is PSI involved?
Is PSII involved?
Where does PSI obtain replacement electrons from?
Where does PSII obtain replacement electrons from?
Is ATP made?
Is reduced NADP made?
8
Chapter 1: Photosynthesis and ATP synthesis
carbon dioxide
(1C)
carboxylation of RuBP
(carbon fixation)
rubisco
ribulose
bisphosphate,
RuBP (5C) intermediate
ADP (6C)
regeneration
of RuBP by Calvin cycle
phosphorylation
glycerate
ATP 3-phosphate,
triose phosphate GP (3C)
(3C)
This is used to triose
make glucose, reduction
phosphate of GP
sucrose and other reduced NADP
carbohydrates.
ATP
oxidised NADP ADP + P
i
of photosynthesis does not change. We can say outside, providing the diffusion gradient that keeps
that light saturation has occurred. Some other it moving into the leaf.
factor, such as the availability of carbon dioxide Carbon dioxide concentration is often a limiting
or the quantity of chlorophyll in the plants leaves, factor for photosynthesis. If we give plants extra
is preventing the rate of photosynthesis from carbon dioxide, they can photosynthesise faster.
continuing to increase. Figure 1.14 shows the relationship between carbon
This relationship is shown in Figure 1.13. Over dioxide concentration and rate of photosynthesis.
the first part of the curve, we can see that rate Figure 1.15 shows the effect of carbon dioxide at
of photosynthesis does indeed increase as light different light intensities.
intensity increases. For these light intensities, light
is a limiting factor. The light intensity is limiting
the rate of photosynthesis. If we give the plant
more light, then it will photosynthesise faster.
Rate of photosynthesis
But, from point X onwards, increasing the
light intensity has no effect on the rate of
photosynthesis. Along this part of the curve,
light is no longer a limiting factor. Something
else is. It is most likely to be the carbon dioxide
concentration.
0
Carbon dioxide concentration
X
Figure 1.14 The effect of carbon dioxide on rate
Light is not a of photosynthesis.
Rate of photosynthesis
limiting factor.
at high light intensity
Rate of photosynthesis
0
Light intensity
0
Carbon dioxide concentration
Figure 1.13 The effect of light intensity on the
rate of photosynthesis. Figure 1.15 The effect of carbon dioxide
concentration on the rate of photosynthesis at
Carbon dioxide concentration different light intensities.
The concentration of carbon dioxide in the air
is very low, only about 0.04%. Yet this substance SAQ
is needed for the formation of every organic 5 a Over which part of the curve in Figure 1.14
molecule inside every living thing on Earth. is carbon dioxide a limiting factor for
Plants absorb carbon dioxide into their leaves photosynthesis?
by diffusion through the stomata. During daylight, b Suggest why the curve flattens out at high
carbon dioxide is used in the Calvin cycle in the levels of CO2.
chloroplasts, so the concentration of carbon
dioxide inside the leaf is even lower than in the air
10
Chapter 1: Photosynthesis and ATP synthesis
GP
TP
0 1 2 3 4 5 6 7 8
Time / minutes
Figure 1.16 The effect of light and dark on the relative levels of TP and GP in a chloroplast.
11
Chapter 1: Photosynthesis and ATP synthesis
SAQ
6 a Explain why the Calvin cycle stops running 7 What effect would you expect a rise or a fall in
when there is no light and the TP is used up. temperature to have on the relative levels of GP,
b Make a copy of Figure 1.16. Add another TP and RuBP? (Assume that the temperature
line to show what you would expect to does not go high enough to denature enzymes.)
happen to the levels of RuBP during this Explain your reasoning.
eight-minute period.
Limiting factors and crop But photosynthesis is not the only process
production which may have limiting factors that affect crop
If one factor is rate limiting, increasing the supply production. Water and each mineral nutrient
of any other factor has no effect on the rate of required by a plant can also limit the rate of
photosynthesis and therefore no effect on crop growth. If the supply of, for example, potassium is
production. On the other hand, increasing the limited in the soil, potassium may be rate limiting
supply of the limiting factor will have a dramatic for crop growth. So, increasing the light intensity
impact on crop production. will have no or little effect on crop production in
For photosynthesis, light or carbon dioxide these circumstances.
could be rate limiting at temperatures within the Analysis of the climate, soil and atmospheric
range tolerated by the crop. However, a farmer environment of a crop can reveal which factor
has limited capacity to affect light or carbon is going to be rate limiting in a range of
dioxide levels in order to increase the rate of circumstances and therefore which factor could
photosynthesis, unless a crop is being grown in a be increased to make maximum impact on crop
specialised glasshouse. production at any one time.
Biofuels
The ability of plants to transfer light energy into that fuel to points of sale, then there is still a net
chemical energy means that they can be used emission of carbon dioxide to the atmosphere.
to provide fuels for us to use for example, for But the greatest problem is the effect that the
generating electricity or in vehicle engines. As increasing quantity of crops to produce biofuels
stocks of fossil fuels run down, and as carbon is having on the availability and price of food. For
dioxide levels in the atmosphere continue to example, as huge areas of land in the USA are
increase, there has been a sharp increase in the use taken over to grow corn (maize) for fuel, there is
of crop plants to produce fuels rather than food. less maize on sale for cattle feed or human food.
For example, rape seed is being used to produce Prices have increased, in some cases so much so
biodiesel, rather than food for animals or humans. that poorer people, especially in neighbouring
At first sight, this would appear to be very countries like Mexico, are finding it much more
good for the environment. Using plants to provide difficult to buy enough food for their needs.
fuels is theoretically carbon-neutral. The carbon We also need to consider effects on ecosystems.
dioxide that is given out when the fuels are Producing large quantities of biofuels will take up
burnt is matched by the carbon dioxide that the large areas of land. There is a danger that some
plants take in as they photosynthesise and grow. countries will cut down forests to provide extra
However, if we take into account the energy that land for this purpose, damaging habitats and
is used in harvesting the plants, converting the endangering species that live there.
biomass to a useful form of fuel and transporting
12
Chapter 1: Photosynthesis and ATP synthesis
Summary
Photosynthesis uses light energy to cause carbon dioxide and water to react to produce carbohydrates
and oxygen. The light energy is transformed to chemical energy in the carbohydrates.
Palisade cells contain large numbers of chloroplasts. They are tall and thin, so light does not have to
pass through too many cell walls before reaching a chloroplast. The chloroplasts are arranged in the
cytoplasm around the outer edge of the cell, and can be moved to maximise light absorption.
Chloroplasts are surrounded by an envelope enclosing the stroma. Inside the chloroplast, membranes
form fluid-filled sacs called thylakoids, which in turn form stacks called grana. Chlorophyll molecules
are embedded in the membranes. The chlorophyll molecules are found within photosystems I and II.
The light-dependent stage of photosynthesis takes place in the thylakoids. Light energy raises
the energy level of an electron in a chlorophyll molecule. The electron is passed down a series of
carriers, generating ATP by photophosphorylation. If only PSI is involved, the electron is returned
to PSI and this is called cyclic photophosphorylation. If the electron is passed to PSII, then reduced
NADP is also produced, using hydrogen ions from the splitting of water. This is called non-cyclic
photophosphorylation.
The light-independent stage of photosynthesis is also known as the Calvin cycle, and takes place in
the stroma. The enzyme rubisco catalyses the reaction of carbon dioxide with ribulose bisphosphate
(RuBP) to form two molecules of glycerate 3-phosphate (GP). This reacts with reduced NADP and
ATP to produce triose phosphate (TP). Most TP is reconverted to RuBP, but one sixth is used to
make carbohydrates such as glucose, sucrose or starch.
Light intensity, carbon dioxide concentration and temperature all affect the rate at which
photosynthesis can take place. The factor that is in the shortest supply at any one time is known as
the limiting factor. If crops are grown in glasshouses, these factors can be adjusted to increase the
rate of photosynthesis and therefore the productivity of the crop.
13
Chapter 1: Photosynthesis and ATP synthesis
Questions
Multiple choice questions
1 The micrograph represents a cross-section of a dicotyledonous leaf. Which region is the main site
of light absorption?
II
III
IV
A I
B II
C III
D IV
2 Where do the light-independent reactions of photosynthesis occur in the chloroplast?
A thylakoid membranes
B stroma
C grana
D matrix
3 The main photosynthetic pigments found in the reaction centre of photosystem II is or are:
A carotenoids.
B xanthophylls.
C chlorophyll a.
D chlorophyll b.
4 In which part of a chloroplast is light energy transferred to chemical energy?
A envelope
B granum
C stroma
D starch grain
continued ...
14
Chapter 1: Photosynthesis and ATP synthesis
5 What are the steps numbered 14 in the diagram of the Calvin cycle shown below?
carbon dioxide
rubisco
ribulose step 1
bisphosphate
intermediate
ADP
glycerate
ATP 3-phosphate
triose phosphate
glucose, step 4
sucrose triose step 2
and other phosphate
reduced NADP
carbohydrates
ATP
oxidised NADP ADP + P
i
6 The electrons lost from chlorophyll a in the reaction centre in photosystem II are replaced by
electrons from:
A reduced NADP.
B photosystem I.
C hydroxyl ions from water.
D hydrogen ions from water.
7 The products of non-cyclic photophosphorylation during the light-dependent reactions of
photosynthesis are:
A glucose, reduced NADP and ATP.
B ATP, oxygen and reduced NADP.
C ADP, carbon dioxide and RuBP.
D hydrogen ions, oxygen and ATP.
continued ...
15
Chapter 1: Photosynthesis and ATP synthesis
8 The graph below illustrates the effect of light intensity on the rate of photosynthesis of a
suspension of Chlorella at 150C and 0.4% carbon dioxide. Which of the following is the most
likely explanation for the plateau labelled Y?
Y
Rate of photosynthesis
0
Light intensity
0
continued ...
16
Chapter 1: Photosynthesis and ATP synthesis
Structured questions
11 a The rate of photosynthesis is limited by certain environmental factors known as
limiting factors.
i What do you understand by the term limiting factor? [2 marks]
ii Identify three environmental limiting factors of photosynthesis. [2 marks]
b The apparatus (Audus microburette), shown below, is used to measure the rate
of photosynthesis.
5cm3 syringe
thermometer
0 10 20 30 40 50 60 70 80 90 100
shoot of Elodea
beaker of water
i Describe how the apparatus shown in this diagram can be used to investigate the
effect of different concentrations of carbon dioxide on the rate of photosynthesis. [4 marks]
ii Describe two precautions which should be taken in setting up and conducting the
experiment to ensure that the results obtained are valid measures of the rate of
photosynthesis under the given conditions. [2 marks]
continued ...
17
Chapter 1: Photosynthesis and ATP synthesis
c The graph below shows how the rate of photosynthesis varies with light intensity at
different concentrations of carbon dioxide.
240
B
200
0.14% CO2 at 20C
160
120 A
40
0
0 1 2 3 4 5 6 7
Light intensity / arbitrary units
H+
1 reduced
NADP
NADP
light H+ light ADP ATP
A B
2e
thylakoid 2e
membrane C
3 4
2 H+
H2O 2H + /2O2
+ 1 H+
continued ...
18
Chapter 1: Photosynthesis and ATP synthesis
a Copy the diagram. Write the labels stroma and thylakoid lumen in the
appropriate boxes labelled 1 and 2 in the diagram. In each case, give a reason
for your answer. [4 marks]
b A and B represent photosystems.
i What do you understand by the term photosystem?
ii Write PSI and PSII in the appropriate boxes labelled 3 and 4.
iii Make a labelled drawing to show the generalised structure of a photosystem. [5 marks]
c Describe the photoactivation of chlorophyll. [3 marks]
d What type of molecule is represented by label C? [1 mark]
e Explain the function of the proton (hydrogen ion) gradient which is produced across the
thylakoid membrane. [3 marks]
f Identify the type of photophosphorylation shown on the diagram. Give a reason
for your answer. [2 marks]
carbon dioxide
enzyme X
ribulose step 1
bisphosphate
intermediate
ADP
glycerate
ATP 3-phosphate
triose phosphate
glucose, step 4
sucrose triose step 2
and other phosphate reduced NADP
oxidised NADP
carbohydrates ADP + Pi ATP
a i State where in the chloroplast the Calvin cycle takes place. [1 mark]
ii The light-independent reactions of the Calvin cycle occur in steps. Copy the
diagram above and identify steps 1 to 4. [4 marks]
iii Name the enzyme X which catalyses the reaction of carbon dioxide with
the carbon dioxide acceptor, RuBP. [1 mark]
iv Name two products of the light-dependent reactions of photosynthesis that are
needed in the Calvin cycle. [2 marks]
v Identify on the diagram the stages where the chemicals identified in iv are used in
the Calvin cycle. [3 marks]
b Gramoxone is a widely used weed killer in the Caribbean. It prevents the
light-dependent stage of photosynthesis from occurring. Explain how it causes
the weeds to die. [4 marks]
continued ...
19
Chapter 1: Photosynthesis and ATP synthesis
Essay questions
14 a By means of annotated diagrams only, describe the structure and functions in
photosynthesis of:
i the external structure of the dicotyledonous leaf. [3 marks]
i a palisade cell. [3 marks]
iii a chloroplast. [5 marks]
b Explain what is meant by the term photosynthetic pigment and explain the difference
between a primary pigment and an accessory pigment. [4 marks]
15 a Describe how light energy is converted into chemical energy in the light-dependent
stage of photosynthesis. [7 marks]
b Explain the main stages of the Calvin cycle. [8 marks]
16 a i Define the term photosynthesis. [2 marks]
ii Photosynthesis is represented by the following equation:
6CO2 + 6H2O C6H12O6 + 6O2
Give four reasons why this equation does not fully represent photosynthesis. [4 marks]
b Discuss the roles of the following in photosynthesis:
i light
ii water
iii photosystems [9 marks]
20